An Overview of The Los Amigos Watershed, Madre de Dios, Southeastern Peru

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An overview of the Los Amigos watershed, Madre de Dios, southeastern Peru
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An overview of the Los Amigos watershed,

Madre de Dios, southeastern Peru
February 2010
Nigel C. A. Pitman, Amazon Conservation Association, Third Floor, 1731 Connecticut

Ave. NW, Washington, DC 20009 USA; npitman@amazonconservation.org
Table of contents
(gray text indicates incomplete sections)
Introduction
Geography and hydrology
Regional context
Geographic locations of the watershed, concession, and field stations
Hydrology
Geology, topography, and soils
Regional context
Local surface geology
Landslides and other landscape disturbances
Topography
Soils
Climate and seasonality
Climate overview
Weather at CICRA, Cocha Cashu, and Puerto Maldonado
Seasonality
Vegetation
Regional overview and recommended bibliographic sources
History of botanical work to date at Los Amigos
The flora of Los Amigos
Vegetation of the Los Amigos watershed
Vegetation types in the Los Amigos watershed
Forests around CICRA
Fauna
Composition, structure, and dynamics
Diversity and endemism
Rare and threatened species
Human history
Introduction
Deep history
1900-1982
The mining camp (1982-1999)
The logging boom (1999-2002)
The age of ACCA (2000-present)
Uncontacted indigenous groups
Research in Los Amigos
Research before ACCA
The first six years (2000-2006)
Research priorities
Acknowledgments
Annotated bibliography
1
An overview of the Los Amigos watershed, uncorrected draft, February 2010
Appendix 1. A list of recommended texts for biologists working in Madre de Dios

Appendix 2. Research projects active at Los Amigos, 2000-2006
Appendix 3. Companion notes for the principal trails at the Los Amigos Research Station
Appendix 4. A preliminary red list for Los Amigos
Appendix 5. Los Amigos in numbers
Appendix 6. Public health at Los Amigos
2
INTRODUCTION
“...and ever since some maps of South America have shown a short heavy line
running eastward beyond the Andes, a river without a beginning and without an
end, and labeled it the River of the Mother of God. That short heavy line... has
always seemed the perfect symbol of the Unknown Places of the earth.”
-Aldo Leopold (1924)

“The River of the Mother of God”
Nearly a century has passed since Leopold wrote those lines, and southeastern Peru is
no longer one of Earth’s Unknown Places. Although most of it remains a roadless
wilderness, and vast stretches will remain unexplored for decades to come, its flora and
fauna are very well known relative to other areas in the Amazon basin. Most of the
region’s vascular plants and vertebrates have been collected and identified, more than
2,200 articles, books, academic theses, and reports have been written about the biology
of the region (Pitman et al. 2007), and at least nine biological stations are currently
active in Madre de Dios. Southeastern Peru is also one of the best-known corners of
Amazonia for non-scientists. Over the last two decades it has replaced Iquitos as Peru’s
leading destination for Amazonian ecotourists—some 70,000 visit each year (Goulding
et al. 2003)—and its national parks are world-famous.
Thus it is surprising how hard it remains to track down basic information on this
landscape: how rainfall varies across the region, which soils are dominant, or which
direction the wind blows in April. Part of the reason is that while a great deal has been
written about southeastern Peru, most publications are either highly specialized articles
written in English and published in scientific journals in the United States or Europe, or
reports, theses, and assorted gray literature written in Spanish, published or distributed
privately in Peru, and available in a few scattered libraries in Lima, Cusco, and Puerto
Maldonado (Pitman et al. 2007).
This document presents basic information on the geography, biology, and history of the
Los Amigos watershed in southeastern Peru. It is intended as a background for research
in and around the Los Amigos field stations. I have tried to include enough specialized
information to make it a useful source of facts on the region without entering into great
detail on any one topic. Earlier descriptions of the Los Amigos landscape are given by
ACCA (2001b) and Forsyth et al. (2002). Valuable descriptions of other sites in Madre
de Dios are given by Terborgh (1983) for Cocha Cashu, Pitman (2000) for Manu, Erwin
(1985) and Napravnik Pesce (2004) for Tambopata, and ONERN (1965; 1972; 1977;
1982) for a variety of other sites around Madre de Dios. A broader overview of the
Madre de Dios watershed is given by Goulding et al. (2003) and a survey of Amazonian
Peru has been published by Kalliola et al. (1993); both are highly recommended.
Appendix 1 of this document lists especially helpful documents for understanding the
landscape, history, and flora and fauna of Madre de Dios.
This survey draws on published and unpublished reports from the region, conversations
with long-time residents of the Los Amigos area, and research to date at Los Amigos.
Since that research remains in the early stages, some of the text here has been adapted
from a preliminary survey of Manu National Park (Pitman 2000).
3
For the moment this document is an incomplete, unpublished draft; corrections and other
suggestions are welcome and should be emailed to the author. To cite this version of the
manuscript, please see the instructions in the acknowledgments section.
GEOGRAPHY AND HYDROLOGY

Regional context
At the southwestern margin of the Amazon basin, Andean rivers descending to the
lowlands are diverted to the north or to the south by the Fitzcarrald Arch, a low geologic
formation jutting out into the lowlands. Rivers to the north and west of the arch flow into
the shallow Ucayali basin, heading northwards to Iquitos before draining into the
Amazon and continuing eastwards towards the Atlantic. Rivers to the south and east of
the arch flow into the shallow Madre de Dios-Beni basin, skirting the base of the Andes
eastwards before heading north into the Madeira River and finally into the Amazon near
Manaus, ~1,600 air km away (Kalliola and Puhakka 1993).
The Los Amigos River forms part of

this river system to the south of the
Fitzcarrald Arch (Figure 1). At the
scale of the entire Madre de Dios-
Beni-Madeira watershed, the Los
Amigos is a very small river indeed.
Its watershed of 4,415 km2
represents less than half of one
percent of the entire drainage; think
of a twig on a towering kapok tree.
The Los Amigos is the sixth largest
watershed in the Madre de Dios
headwaters in Peru—much smaller
than the Las Piedras (the largest at
nearly 20,000 km2), as well as the
Manu, Alto Madre de Dios,
Inambari, and Tambopata, but
larger than the Colorado, Azul, and
Blanco (Goulding et al. 2003). The
mouth of the Los Amigos lies just
Figure 1. Location of the Los Amigos watershed in 500 km from the beaches of the
western South America. The lake in the lower right- Pacific Ocean, but roughly 2,600
hand corner is Titicaca. Image courtesy of William
Bowen.
km in a straight line from the mouth
of the Amazon.
The Los Amigos watershed is part of the 85,183 km2 department of Madre de Dios.
Madre de Dios is the third-largest Amazonian department in Peru, after Loreto and
Ucayali; 82% of it is below 500 m elevation. The Los Amigos watershed is centrally
located in Madre de Dios, just a few kilometers south of the department’s geographic
centroid. (The centroid, an otherwise trivial cartographic feature, has attained modest
importance in recent years because some databases of biological collections use it as a
default locality for collections that were made in the department but whose labels lack
specific locality data; the irony is that the centroid is in a very remote part of the Las
Piedras watershed where it is unlikely that any biologist will ever collect a specimen.)
The western portion of the Los Amigos drainage belongs to the Distrito and Provincia de
4
Manu, while the eastern portion belongs to the Distrito de Laberinto and the Provincia de
Tambopata.
The Los Amigos watershed is entirely lowland, although both its mouth and its
headwaters are only 50 km north of the Andean foothills. The river runs roughly parallel
to the Andes for its entire length of 353 km.
Geographic locations of the watershed, concession, and field stations

Los Amigos is at about the same latitude as Lima. The same line runs through Salvador
(Brazil), northernmost Madagascar, and northern Australia. (Comparable latitude in the
northern hemisphere runs through Nicaragua, Senegal, and southern India.) Longitude
at Los Amigos is the same as that in Boston (USA) and Tierra del Fuego.
The Los Amigos conservation concession covers 145,918 ha in the lower third of the
watershed, including its southernmost and easternmost points, and has a perimeter of
450 km. (At its establishment the concession measured 135,832 ha and included some
isolated patches that were still logging concessions; those became part of the
conservation concession in early 2005.)
The CICRA station is at approximately 12°34’07”S 70°05’57” W, or 380500E 8610297N

19S in UTM coordinates; elevation is ~268 masl. Elevation on the CICRA terrace is ~41
m above the arbitrary zero mark of the Madre de Dios River at the CICRA dock (see
hydrology) and ~80 m above the city of Puerto Maldonado, but nearly 100 m lower than
the upland terraces around Cocha Cashu. Elevation througout the Los Amigos
watershed is much higher than most of northern Amazonian Peru; Iquitos is at ~100
masl.
The CM1 station is at approximately 12°34’17”S 70°04’29”W, or 383789E 8609498N

19S in UTM; elevation there is ~238 masl. The CM2 station is at approximately
12°26’57”S 70°15’06”W, or 363960E 8623430N 19S in UTM; elevation there is ~259
masl.
The geographic position data given above are taken with GPS near the center of each
station and are accurate to a few tens of meters. There are not yet any permanent
reference markers in the Los Amigos watershed that have been surveyed for geographic
position and elevation with cm-level accuracy; establishing such markers at both CICRA
and CM2 is a short-term goal. In 2003-2004 ACA placed permanent cement markers
along the border of the conservation concession, but precise (i.e., <10 m accuracy)
coordinates and elevations are not yet available for them.
Some other organizations have also placed permanent cement markers in the Los
Amigos area. Markers at the corners of mining concessions are topped with a circular
brass marker inscribed with a place name, a number, and the words “Catastro Minero
Nacional MEM (Ministerio de Energía y Minas).” Coordinate and elevation data for these
markers are available from the Ministerio de Energía y Minas office in Puerto
Maldonado, but it is not known how accurate the data are. The Proyecto Especial de
Titulación de Tierras (PETT) of the Ministerio de Agricultura has also placed some
cement markers in the region to demarcate private properties. One is located in the
CICRA clearing, near the weather station, and reads “PETT-CBS-6”. The geographic
information associated with the PETT markers is not especially reliable (D. Pogois,
personal communication).
5
A variety of cartographic resources are available for the Los Amigos watershed.
Topographic maps at the scale 1:100,000 are available in Carta Nacional sheets 25-u,
25-v, and 26-v, for sale in Lima at the Instituto Geográfico Nacional
(www.ignperu.gob.pe). Dozens of GIS layers, satellite images, and maps of the
watershed are now publicly available at atrium.andesamazon.org.
Madre de Dios, like all of Peru, is five hours behind Greenwich mean time; it is in the
same time zone as the eastern seaboard of the United States, but does not follow
daylight savings time. At the December solstice, the sun in this part of Peru rises at 5:09
and sets at 6:01. At the June solstice, the sun rises at 5:56 and sets at 5:20. The
shortest day of the year (21 June) has 11:24 of sunlight, while the longest day (21 Dec.)
has an hour and a half more. The sun passes directly overhead Los Amigos on around
27 October and 16 February. Days at Los Amigos are never symmetric around noon,
i.e., there is always more daylight between sunrise and noon than between noon and
sunset. This is because we are significantly east of the Standard Time Meridian for our
time zone (75°W). Another result of this is that the sun always reaches its zenith at Los
Amigos before 12:00 noon local time.
The declination in Los Amigos, as of June 2006, is 5°16’ W, increasing by 0°9’ W each
year. In other words, compasses at Los Amigos point at a magnetic north pole that is
five degrees to the west of the line to the geographic north pole. To walk in a straight line
towards the geographic north pole with a compass at Los Amigos, one must aim five
degrees east of magnetic north. In general, to walk at n degrees to geographic north,
one must follow the compass n + 5 degrees. Declinations change over time; the current
declination for Los Amigos can be calculated at the following website:
www.ngdc.noaa.gov/seg/geomag/jsp/Declination.jsp
The closest settlement to CICRA is Boca Amigos, ~3 km to the SE and about ten
minutes’ travel downriver from the station by outboard canoe. The closest large town is
San Juan Grande, ~12 km to the W of CICRA, about 40 minutes’ canoe travel upriver
from the station. The closest city is Puerto Maldonado, ~90 km E of CICRA, about three
hours’ travel downriver by outboard canoe and an hour’s travel by road. Appendix 5
provides air distances from CICRA to various landmarks and field stations in this part of
the world.
Hydrology
Large rivers
Very little of the physical setting in the Los Amigos watershed makes sense without
reference to the river systems that successively build up and tear down the landscape.
There are at least two kinds of large rivers in the area: steep-gradient, high-energy
torrents roaring down from the Andes—like the Alto Madre de Dios and the Inambari—
and lower-gradient, meandering lowland rivers like the Los Amigos, the Manu, and the
Las Piedras. The high-gradient rivers tend to produce braided channels and carry a
heavy sediment load—from sand to gravel to large boulders—while the lowland rivers
twist back and forth across broad floodplains dotted with abandoned oxbow lakes,
moving mostly silt and sand. High-gradient rivers are generally classified as white-water
rivers, with neutral chemistry and high levels of suspended sediments. Lowland rivers
throughout Amazonia tend to be slightly more acidic and with a lighter sediment load,
and are placed at an intermediate position in the black- and white-water dichotomy
6
(Kalliola and Puhakka 1993). In contrast with other areas of Amazonian Peru, there are
no large blackwater rivers in Madre de Dios.
Rates of lateral migration are poorly known for the high-energy rivers, and course
changes may be sudden rather than gradual. As the beds of these rivers accumulate
more and more gravel, rocks, and other debris, they gain elevation relative to the
surrounding landscape, thereby increasing the likelihood that rivers will find a lower
gradient elsewhere and jump abruptly to new routes during high water events. Thus it is
no surprise that the soils of the high terraces to either side of the Alto Madre de Dios
River (and many other locations in the uplands south of the Madre de Dios) are full of
gravel; they are the product of high-energy braided rivers that have scythed back and
forth across the landscape for some ten million years, leaving trails of rocky debris from
the Andes.
Meandering rivers such as the Manu carry mostly fine particles, and migrate laterally
through a sandy floodplain at a rate of 5-25(-200) m/yr. It has been estimated that such
rivers can cut down and rebuild their several-kilometer-wide floodplains every few
hundred years (Terborgh 1990; Kalliola and Puhakka 1993). However, the ubiquity of
oxbow lakes and their sometimes impressive age—Cocha Cashu, for instance, may be
~2,500 years old (M. Silman, personal communication)—suggest that some of the
floodplain has been untouched by river migration for several thousand years (but see
Foster 1990). Schroeder and Zimmermann (unpublished data) have estimated from a
comparison of remote sensing images taken of the Manu River in 1962 and 2001 that
floodplain 1 km distant from the river is disturbed, on average, every 2,000 years by
meandering dynamics. Puhakka and Kalliola (1995) have quantified some aspects of
river meandering and oxbow lake formation on the Madre de Dios and Los Amigos
rivers. The oldest known aerial images of the landscape around Los Amigos were taken
in 1960, so it will soon be possible to reconstruct a more complete meander history of
both rivers.
Flooding events on all rivers in the region are poorly recorded. Since September 2001
the level of the Madre de Dios River has been recorded twice daily (morning and
evening) at the CICRA docks. River level is measured against marked posts set out
around an arbitrary zero. The +5 m level is maintained, for the time being, as a mark on
the supports of the CICRA sign at the docks; a more permanent calibration point is
needed, as the riverbank beneath the sign appears to be slumping. The best
measurements to date suggest that the arbitrary zero on the CICRA river gauge is at
approximately 227 masl. A similar river gauge was established on the Amigos River at
the CM2 station in December 2005 and is likewise being monitored twice daily. River
level is also being monitored on the Madre de Dios at Puerto Maldonado (C. Cañas,
personal communication), but the Los Amigos gauges and the Puerto Maldonado gauge
have not been calibrated one to the other.
Data from the CICRA river level gauge are given in Figure 2. Low water is typically in
September, while high water is typically between December and February. The
difference between the highest record and the lowest record in the dataset is 9.47 m,
and the river is capable of rising or falling as much as 3 m in a 24-hour period.
Significant areas of the CICRA trail system in the floodplains become impassable during
high water on the Amigos and Madre de Dios rivers, but we do not yet have an effective
way to relate river level as measured at the CICRA gauge with the flooding depth or
duration of floodplain and swamp forests around the station.
7
Figure 2. Variation in the level of the Madre de Dios River from August 2001 to April 2006, as
measured twice daily at the Los Amigos dock. The horizontal lines mark the arbitrary zero. The
absolute maximum and minimum values are marked by asterisks. The highest river level (+5.62
m) was recorded on 21 January 2003, when a state of emergency was declared in the region
because of severe flooding. The lowest river level (-3.85 m) was recorded on 9 September 2005,
at the end of one of the driest dry seasons ever recorded in Amazonia. The inset (red lines)
shows all five years on the same vertical axis.
Barthem et al. (2003) summarize in detail the various changes in water chemistry and
other river characteristics between periods of high and low water. During low water, the
Madre de Dios is slower, warmer, more acidic, more transparent, and with higher
conductivity. There is no significant difference in water oxygen levels in the two seasons.
Goulding et al. (2003) speculate that approximately half of the water in the Madre de
Dios River is runoff from the Andes; the rest is from the lowlands. Heavy rainfall in the
Colorado watershed (the next large tributary of the Madre de Dios upriver from the Los
Amigos) turns the Madre de Dios a characteristic bright orange, due to erosion of the
colored clay that gives the Colorado its name. Water in the Los Amigos river is warmer
than the Madre de Dios,
The channel of the Madre de Dios is ~xxx m wide at the CICRA dock. Goulding et al.
(2003) give maximum depths of 15 m for the Madre de Dios, with high water means of 7-
10 m and low water means of 2-5 m, and high water velocities of 11-14 km/hour.
Lakes
Nearly all lakes in the region are oxbows—known locally by the Quechua word for lake,
cocha—sections of channel that have been abandoned by the meandering river and
8
begun a new life as lakes in the floodplain. There are ~40 cochas in the Los Amigos
concession, and several larger ones in the Madre de Dios floodplain around the mouth
of the Los Amigos. The only significant lake in the region that does not appear to be an
oxbow is the Pozo Don Pedro, a tiny blackwater lagoon in the palm swamp to the
northwest of CICRA. Mining operations have also created dozens of small ponds in the
Madre de Dios floodplains around the station; it is not clear how long these will persist.
Cocha water undergoes seasonal changes similar to those seen in large rivers
(Goulding et al. 2003). Although most cochas are no longer connected to the river that
formed them, and receive river water only during especially high floods, most do receive
water year-round from small streams that drain the uplands. As a result, water acidity
and conductivity in cochas tend to be intermediate between that of rivers and upland
streams. Cochas vary significantly in age and size, for obvious reasons, but also in water
chemistry, aquatic plant communities, and fish communities, for reasons that are not well
understood (Davenport 2003).
Streams
Most rain in Madre de Dios falls in the uplands, far from lakes and rivers. Some of what
does not evaporate back into the atmosphere runs directly off the uplands via small
streams. Other rainwater is absorbed by the soil and filtered down to a water table well
above river level, from which it seeps laterally out of hillslopes as small springs,
supplying the streams during periods of low rainfall. Water in these streams is typically
cooler, more acidic, and poorer in nutrients than that in large rivers (Goulding et al.
2003). Streams in the uplands often trace the dividing line between soils of different
textures, so that one bank of a stream may be predominantly sandy while the opposite
bank is mostly clayey (Osher & Buol 1997).
All streams, seeps, swamps, and other bodies of water on the CICRA trail system were
visited and mapped, and their water characterized chemically in May 2006. An
annotated map should soon be available (G. Barbieri, personal communication).
Drainage pattern dendritic (Khanal 2006).
GEOLOGY, TOPOGRAPHY AND SOILS

Regional context
Los Amigos sits in a shallow depression at the southernmost end of the crescent-shaped
Amazonian foreland basin that runs along the eastern base of the Andes. The Madre de
Dios-Beni sub-basin south of the Fitzcarrald Arch is in an aggradational stage, i.e., it is
still accumulating alluvium carried down from the Andes by high-energy, high-volume
rivers like the Alto Madre de Dios and the Inambari.
The geologic parent material underlying the region is recently deposited alluvium of the
same kind. All of the upland terraces aged to date in the region date to <200,000 years
old, with some dates possibly as recent as 30,000 years old (see next section). The
material is anything but homogeneous. Stratigraphic sections reveal a jumble of
differently-textured material corresponding to different kinds of historical deposition
events—gravel layers indicating ancient river bottoms, sand deposited in point bars by
laterally migrating rivers, and silt and clay deposited by occasional flooding events
(Räsänen 1993).
9
Local geology
Information about the geology of the Los Amigos concession comes from: 1) seismic
profiles of the oil industry (Huertas Castillo 2002), as well as at least two exploratory
wells dug in the Los Amigos watershed (north of the concession) during 19xx by xxx; 2)
the official Peruvian government geological survey of the area, carried out by the
Instituto Geológico Minero y Metalúrgico and published as a book with geological maps
at 1:20,000 (Chávez et al. 1998, available for sale at INGEMMET’s office in Lima); and
3) studies of surficial geology by scattered groups of geologists and paleontologists.
ACA has not yet obtained the relevant information from historical oil exploration in the
region, in part because much of it remains privately owned.
Antoine et al. (2003) show the Los Amigos watershed sitting on top of a 3 to 4-km thick
slab of alluvium deposited during the Neogene, i.e., over the last 25 million years. The
dominant geological formation at the surface in the Los Amigos concession is the Madre
de Dios Formation, dated indirectly at upper Miocene to Pliocene. Antoine et al. (2003)
describe this formation as a series of alluvial, lacustrine, and tidal deposits, and suggest
that some of it was deposited by an inner sea in the region during the upper Miocene.
The Madre de Dios Formation overlies the older but similar Ipururo Formation, which
becomes dominant to the north towards Acre. Below these formations is a basement of
older alluvium. The basement tilts downwards in the direction of the Andes, pushed
down by the weight of that huge mountain range. This same phenomenon is believed to
be uplifting the Los Amigos landscape (Antoine et al. 2003). Presumably this tilting is
also the reason why the Madre de Dios and the Los Amigos rivers do not run north,
directly out into the Amazon lowlands, but instead run parallel to the base of the Andes.
The massive landslide 500 km NW of CICRA, overlooking the Madre de Dios, provides
one of the longest and most accessible stratigraphies in southeastern Peru. Its deep red
alluvial soils, sitting on a grey formation closer to the river, have been described in detail
by Chávez et al. (1998), Antoine et al. (2003), and Hovikoski et al. (2005). The same
outcrop has also produced some interesting plant and animal fossils. Antoine et al.
(2003) carbon-dated an unidentified piece of wood embedded in the hillside a few
meters above the river level, and estimated an age >45,000 yr. Campbell et al. (2000)
and Alberdi et al. (2004) describe a fossil gomphothere discovered in the same
landslide. The mining camp that occupied the CICRA terrace in 1982-1999 had on
display the fossilized skull of a huge crocodilian. It is not clear where the fossil was
discovered, but it is reasonable to suppose that it was also in the vicinity of CICRA. It
seems likely that the skull belonged to Purussaurus brasiliensis, a Miocene giant that
measured 14-18 m long and weighed more than Tyrannosaurus rex.
The youngest geological deposits are the thin strips of active floodplains along the major
rivers. These accumulate sediment every few years, when rivers overflow their banks
onto the adjacent floodplains. These deposits, and the river bottom itself, are the only
places on the landscape with appreciable amounts of gold. Because the gold originates
from the Andean foothills and travels down rocky rivers, most of the gold around Los
Amigos is in the form of fine particles—nuggets are ground to sand on their way down
the rocky rivers. The richest alluvial deposits are downriver of the Río Colorado, which
drains the region’s principal mining area, the Río Huaypetuhue.
Earthquakes, landslides, and other landscape disturbances

Los Amigos is about 50 km north of the Tambopata Fault, but earthquakes are rare in
the region. There is also no significant vulcanism, past or present, in the region. Some
10
small hot springs exist along the upper Alto Madre de Dios and are being developed for
tourism (C. Cañas, personal communication).
Probably the most important large-scale physiographic disturbances in the region, apart
from the meandering rivers (see above), are landslides. These may be triggered by a
combination of tremors, treefalls, and massive rainfall, or they may be the culmination of
years of minute slippage. Larsen and Simon (1993), based on observations in Puerto
Rico, suggested that 200 mm of rainfall in a 72-hour period is likely to set off a large
number of landslides in hilly tropical terrain. This threshold has been reached once at
CICRA in six years (240 mm).
Khanal (2006), examining 2003 air photos of a 1,634-km2 area in the lower Los Amigos
watershed, mapped 381 landslides. The largest measured 4 ha; the mean size was 0.24
ha. Together, these landslides accounted for 0.5% of the study area. Landslides were
spatially autocorrelated up to around 2 km, but they were not clumped where one might
naively expect them to be: around the steep cliffs overlooking the floodplain of the Los
Amigos River. Instead, they seem to cluster in places where the terraced uplands and
hilly uplands border each other, and where the steep cliffs along the river have long
since collapsed to form young watersheds (N. Pitman, personal observation).
In 2005 Pascual Flores recalled the three major landslides that occurred during the
nearly 20 years that he has lived on the CICRA terrace. None of them took place close
to CICRA; all of them were in the bare hillside a few hundred meters upriver from the
CICRA docks. Flores saw one of the landslides with his own eyes, and remembers being
unimpressed by the spectacle until the cascading debris hit the river, whereupon it threw
up a massive wave that traveled a long ways up and down the Madre de Dios River and
washed into the forest on the opposite bank. Flores tells of walking through the forest on
the opposite bank shortly after the landslide and finding several large fish on the forest
floor, stranded there by the wave.
Physiography
Perhaps in compensation for the remarkable paucity of information on the region’s soils,
hydrology, geology, and climate, geographers have described physiographic features of
southeastern Peru in exhaustive detail. Part of the reason is that describing topographic
features is relatively easy to do with aerial photos or satellite images. Another reason is
that many biogeographers have hoped that physiographic units might serve as proxies
for plant and animal distributions: indeed, the leading forestry maps of Amazonian Peru
are still based directly on topography rather than vegetation. Thus one description of the
Los Amigos watershed distinguishes ten different physiographic units, encompassing
terraces and hills of varying degrees of dissection, elevation, and steepness (ACCA
2001; the units seem to have been determined from a physiographic map of the region,
but the original source is not given in the report).
Foster (2001) sensibly lumped these into three units which form fairly discrete and
obvious blocks in Landsat images of the Los Amigos region: floodplains, terraced
uplands, and hilly uplands. The terraces, “notable for being very flat with only infrequent
cutting by small streams.... are the very western tip of a formation that forms a broad
regional arc of weakly dissected uplands. This formation does not go south of the Rio
Madre de Dios except in the vicinity of Puerto Maldonado (where it crosses over to just
beyond the Rio Tambopata), but instead sweeps northeast into Pando, Bolivia, eastern
Acre, Brazil, and beyond” (Foster 2001).
11
The hilly terra firme, he writes, are “highly dissected steep hills ~50-100 m high or
higher. This is the southernmost end of a large regional physiographic formation,
interrupted only by rivers, that stretches northwest and north for hundreds of kilometers
into the Ucayali Department of Peru, western Acre, Brazil, and beyond. It also does not
pass south of the Rio Madre de Dios, though it does appear to be on both sides of the
Manu floodplain above the Rio Pinquen.”
Foster (2001) offers a variety of potential explanations for the existence of two
physiographically distinct kinds of uplands. The best-supported seems to be that hills
and terraces reflect the results of weathering on different soil types. As Osher and Buol
(1997) have noted near Puerto Maldonado, coarse soils tend to produce gradual slopes,
while fine-textured soils tend to produce steeper, convex slopes. Since upland terraces
in Madre de Dios contain significant amounts of sand, the prediction is that soils in the
hilly uplands have higher concentrations of clay.
(Availability of topographic data for the region.) Topography of the Los Amigos region is
given in Figure x. The highest elevation within the watershed is ~433 m, and the lowest
elevation is ~226 m (Khanal 2006). Terraced uplands and floodplains are the dominant
physiographic formations around CICRA. All three main formations are accessible from
the CM2 station.
Soils
Soils in Madre de Dios are either very complex or very simple, depending on one’s
perspective. On the one hand, since the parent material that modern soils are derived
from is a jumbled mosaic of clayey materials, sandy materials, silty materials, gravel,
and various mixtures of these deposited by thousands of rivers that meandered
unpredictably across the ancient landscape, soil texture can vary dramatically in the
course of a short walk through a single forest type. For example, soils collected at half a
dozen upland sites between Cocha Cashu and Los Amigos had sand contents ranging
from 27 to 85% (N. Pitman, unpublished data). On the other hand, despite this significant
spatial heterogeneity in texture, the vast majority of upland soils in the region fall into just
two classes in the soil taxonomy system: the Ultisols and Inceptisols that dominate much
of western Amazonia (Linna 1993, Mazer 1996, Osher & Buol 1997, Phillips et al. 2003,
Pitman et al. 2001, Riley 1994).
It is still too early to draw detailed conclusions about the fertility of Madre de Dios soils
relative to other sites in Amazonia or the Neotropics, but in general they occupy the
fertile end of the gradient. There is no doubt that floodplain soils in the region are fertile;
because much early work was done on the Cocha Cashu floodplain, the region gained a
reputation among tropical biologists for relatively rich soils. Pitman et al.’s (2001)
comparison of upland soils in Madre de Dios with the fertile upland soils of eastern
Ecuador seemed to confirm this impression; no significant differences in nutrient
availability were found.
The most detailed study of upland soils in Madre de Dios to date is an examination of a
forested site on the road between Puerto Maldonado and Laberinto (Osher & Buol
1997). They found that “all upland soils classify as Ultisols, in clayey, fine-loamy or
coarse-loamy families.” Upland soils were very acidic, with pH of 3.6-4.7. Acidity
decreased, while clay content increased, with depth. The dominant clay mineral was
kaolinite, while the dominant mineral in sandy samples was quartz.
12
Terra firme soils are consistently sandier, more acidic, and poorer in nutrients than
floodplain soils (Mazer 1996). Among flooded forest samples, extremely young soils (i.e.,
new levees created by river dynamics) tend to be less acidic than older soils, and to
concentrate nutrients at much greater depths than older soils (Riley 1994). The best
study of alluvial soils in Amazonian Peru is from Loreto, but it probably applies well to
floodplains in Madre de Dios (Paredes Arce et al. 1999). Thus a very simplistic picture of
soil diversity in Manu shows >80% of the landscape covered with mostly coarse-textured
and relatively poor upland soils derived from young alluvium and the remaining 10%
composed of thin bands of fine-textured and relatively rich alluvial soils along streams
and rivers.
The most striking edaphic feature in southeastern Peru are the clay licks scattered
across the landscape and visited by birds, mammals, and other animals (Emmons &
Stark 1979). These clay licks, known as colpas in Spanish orthography, ccolpas or
ccollpas in Quechua, are valued by hunters and ecotourists for the abundance of animal
life they attract. Twenty-five years after the first paper on the region’s clay licks (Emmons
and Stark 1979), biologists and soil scientists have yet to pinpoint the reasons that lead
animals to eat soils (Brightsmith & Munoz-Najar 2004). Even less is known about the
distribution of these economically and biologically important features across the
department, across Amazonian Peru, or at larger scales in the Amazon basin. As of
2005, the map of clay licks on the Los Amigos River (Figure 3) was the only colpa map
in existence for Madre de Dios, and perhaps for all of Amazonian Peru (D. Brightsmith,
personal communication).
Figure 3. A preliminary map of clay licks in the Los Amigos watershed.
13
CLIMATE AND SEASONALITY

Climate overview
Climate in the region is warm, wet, and decidedly seasonal, with most rain falling outside
of a pronounced three- to four-month dry season (see earlier overviews in Barthem et al.
2003, Schenck 1999, Terborgh 1983, 1990; Terborgh et al. 1986).
Large-scale weather maps show Los Amigos at the intersection of two climatic gradients
(Kalliola et al. 1993). To the south, mean annual temperatures drop at least 20°C as
elevation rises from <500 m in the lowlands to >4,000 m in the Andes, and along the
same gradient precipitation reaches >7,000 mm at intermediate elevations, before
decreasing again at higher elevations. To the east, rainfall declines dramatically, and
forests give way to savanna in large areas of Bolivia, and Brazil. Figure x shows how the
rainfall and seasonality of Los Amigos and other sites in Madre de Dios compare to well-
known study sites elsewhere in the tropics.
Long-term, high-quality weather data from Los Amigos do not yet exist, and so a picture
of the watershed’s climate must be pieced together from a variety of other sources.
While there is an abundance of scattered data—including a five-year record from CICRA
2000-2006, decades-long data from the five Peruvian government climate stations in the
surrounding lowlands (Puerto Maldonado, Iberia, Iñapari, Pakitza, and Puerto
Esperanza), several years’ worth of data from a variety of privately operated weather
stations (e.g., Cocha Cashu, Tambopata Research Centre, Explorers’ Inn), and the
personal observations of long-term residents and researchers (e.g., Foster and Terborgh
1998)—no one has yet compiled these into a single dataset. Part of the reason is that
the best climate record in the region—from Puerto Maldonado, where rainfall,
temperature, and humidity have been recorded daily (but not continuously) since 1948—
is not publicly available. The Puerto Maldonado data can be purchased from Peru’s
Meteorology and Hydrology Service (SENAMHI) at very high cost (~US$9,000 as of
December 2004). In the meantime, historical summaries (i.e., monthly means) of many
of the SENAMHI stations are available on various public access websites.
Collating these data is a top short-term goal of ACA, because several important
questions about the regional climate remain unanswered. How does climate vary across
the lowlands of the region, over hundreds of kilometers (Cocha Cashu in the west vs.
Cusco Amazónico in the east), over dozens of kilometers (CICRA vs. CM2), over a few
kilometers (CICRA vs. the end of Huangana trail), and between different physiographic
features (high terraces vs. adjacent floodplains)? How have the frequency and intensity
of friajes (cold snaps) and windstorms varied over the 40-year climate record for Madre
de Dios? How does the climate change during El Niño/Southern Oscillation and La Niña
events? How do climatic factors affect plant phenology and animal behavior?
Fortunately, the region’s weather is constant enough in many aspects that a ragged
dataset provides a reasonably good picture of its basic features. Even Terborgh’s (1983)
initial survey of Cocha Cashu’s weather, based on a single year of data recorded in
1976-77, still provides an accurate picture of basic weather patterns, despite several
subsequent years’ worth of record-collection at the same site. Casual observations of
professional ten-day forecasts of the weather in Puerto Maldonado (available online at
www.weather.com, www.accuweather.com, and other such sites) do a reasonably good
job of predicting conditions at Los Amigos.
14
Weather at CICRA, Cocha Cashu, and Puerto Maldonado

(Description of measurement methods and available data from the three sites.)
Rainfall and temperature

Mean annual rainfall at CICRA in 2000-2006 was between 2,700 and 3,000 mm (Table
x). There are at least three different ways to calculate this value from the available data.
Summing monthly means for all 65 months of complete data gives 2,735 ± 853 mm (1
SD). Averaging the annual rainfall in the three complete years of data (2001-2003) gives
2,995 ± 455 mm. Finally, since >80% of rainfall in southeastern Peru falls during the
October-to-May wet season, one can also examine annual rainfall totals for entire wet
seasons, rather than breaking each wet season in two at the December/January mark
and treating half the wet season data as belonging to one year and the other half as
belonging to the next. When 12-month rainfall averages that include entire wet seasons
are examined, averaging the three complete years of data gives 2,989.4 ± 348 mm.
All of these numbers are significantly higher than long-term means from Puerto
Maldonado (2,259 mm) and Cocha Cashu (2,200-2,500 mm), but it is premature to
conclude that these differences are statistically significant, since the CICRA record is
based only on recent years. When annual rainfall at CICRA in 2000-2005 is compared
for the three sites, xxx. In all three datasets rainfall is markedly seasonal, with >80% of
the precipitation falling between October and April, when the convectional clouds of the
intertropical convergence zone visit these latitudes. The driest months are June, July,
and August, each averaging <80 mm of rain in both records, with May and September
averaging only slightly more than 100 mm (Figure x). Relative humidity at Puerto
Maldonado tracks precipitation, with only the seasonally dry months averaging below
80%. At CICRA, relative humidity averaged 87.3% between September 2004 and April
2005. Absolute minimum was 29.2%, recorded in a period of September-October 2004
when relative humidity around noon was often below 40%.
15
600
2000
2001
2002
500
2003
2004
2005
2006
400
Precipitación (mm)
300
200
100
0
ene feb mar abr may jun jul ago set oct nov dic
Figure 4. Monthly rainfall totals at the Los Amigos Biological Station from December 2000 to June
2006.
These average rainfall values mask a great deal of year-to-year variation, both in total
annual rainfall and in the strength and duration of the dry season (Terborgh 1983).
Annual rainfall at CICRA has been as low as 2,612 mm (2001) and as high as 3,498 mm
(2003). Cocha Cashu has ranged from 1,600 mm (1983-84) to 2,700 mm (1993-95), and
Puerto Maldonado from <2,000 to >5,000 mm (Barthem et al. 2003). Much of this
variation appears to be due to a high variance in rainfall in the transitional months
between the wet and dry seasons—April, May, September, and October (Figure 4). At
CICRA, 472 mm of rain fell in April 2003, while <40 mm fell during April 2005. For every
month of the year in the Puerto Maldonado dataset, maximum rainfall recorded for any
given month is more than an order of magnitude higher than the minimum for that same
month.
Rainfall in many tropical climates peaks in the afternoon, after many hours of
evaporation and transpiration, but this is not the case for Los Amigos (Figure x). At Los
Amigos rain is more likely during the morning; 25% of all rainfall occurs between 10 AM
and 1 PM. Rain is least likely between 9 and 10 PM (just 0.7% of the total), and rises
steadily from that minimum value to its peak value between 12 noon and 1 PM (which
accounts for 9.6% of the total). After 1 PM the probability of rain declines steadily
through the afternoon.
16
Figure x. Distribution of rainfall during the day at Los

Amigos, based on data collected half-hourly at
CICRA between September 2004 and October 2007.
Temperature shows a much milder seasonal signal. The dry season is slightly cooler
than the wet season, but monthly means never depart from the range of 21-26° C. The
highest temperature recorded at CICRA since 2000 is 39° C (37.1° C with the automated
station), at Cocha Cashu since 1976 33°C, and at Puerto Maldonado since 1948 40°C.
The first two complete years of data from the automated weather station at CICRA
indicate a very slight increase in average temperature from 2005 to 2006. The mean
temperature for 2006 (24.10° C) was 0.28% higher than that for 2005 (24.03° C).
Friajes
One of the region’s most important climatic features are the periodic cold fronts, known
locally as friajes, that sweep through Madre de Dios during the austral winter. These are
large masses of cold air driven northwards from Patagonia into the Amazon basin—the
southern hemisphere equivalent of the arctic fronts that periodically sweep down into the
continental United States during the boreal winter (Marengo et al. 1984, 1997). During
these phenomena temperatures can drop as much as 20° C in under 24 hours. Most
friajes last between one and four days, but some last for more than 10 days, and
consecutive cold fronts can keep daily minima below 20°C for as long as a month (Table
x, Figure x). The historical lows at CICRA, Cocha Cashu and Puerto Maldonado are 8°
C, 8° C, and 4.5° C respectively.
The arrival of a friaje at Los Amigos is typically heralded by strong winds sweeping in
from the south and west. On the Los Amigos terrace the approach of these winds is
audible several minutes before their arrival, as they come roaring through the
surrounding forest or up the river. The first gust is often the strongest. Accompanied by a
harrowing roar from the trees around camp, it comes tearing into camp, knocking over
equipment, ripping off roof cornices, and sending the contents of clotheslines soaring off
the cliff. For the next 30 minutes to an hour, these winds continue to blow, surging and
subsiding like waves on a roughly two-minute cycle. In most cases the wind is
17
accompanied by heavy rain, which causes additional havoc as it is driven into rooms and
laboratories by the strong winds.
In the forest on the CICRA trail system trees and branches are commonly flung down by
these storms, and witnessing one from inside the forest is a dangerous and nerve-
wracking experience. Although the most violent winds typically blow for less than an
hour, steady westerly and southerly winds often continue for several hours after the front
has passed. At the station, one lasting effect can be seen in the young trees around
camp, many of whose trunks are permanently angled towards the west as the result of
being pressed so insistently in that direction by these winter storms.
Figure x. Frequency of cold fronts, or friajes, at the Los Amigos Biological Station during the
winter months of 2005-2007. Each graph shows temperature data at 30-minute intervals from 1
April to 30 September. In each graph the vertical axis ranges from 10 to 40 degrees Celsius, with
gridlines at 20° and 30° C.
As the wind and rain die down, air temperature drops precipitously, often recording the
cold front's lowest temperature during the first 24 hours after its arrival (Figure x). For the
next several days, under skies that may remain overcast or cloud-free, temperature
maxima and minima typically warm by about 1° C per day until normal temperatures are
again attained or another cold front arrives.
While the frequency, duration and intensity of friajes vary from year to year, data
collected at Los Amigos in the years 2005-2007 show some remarkably stable patterns.
18
To compare patterns among these years, I arbitrarily defined friajes as more than two
consecutive days with daily minima <20° C, and severe friajes as more than two
consecutive days with daily minima <15° C. By those definitions, the number of events
varied very little in the winter months of 2005-2007: 14-17 friajes and 4-5 severe friajes
per winter (Figure x, Table x).
Table x. Frequency of cold fronts, or friajes, at the Los Amigos

Biological Station during the winter months of 2005-2007.
2005 2006 2007

No. friajes (at least two consecutive
14 16 17
days with minima <20° C)
Median no. consecutive days with
4 6 5
minima <20°C
Longest no. consecutive days with
30 17 15
minima <20° C
Proportion 30-minute records <20° C 24.2% 22.8% 24.2%
No. severe friajes (at least two
5 4 5
consecutive days with minima <15° C)
Median no. consecutive days with
3 2.5 3
minima <15°C
Longest no. consecutive days with
4 3 4
minima <15° C
Proportion 30-minute records <15° C 4.3% 2.0% 4.2%
It is not yet clear what effect El Niño/Southern Oscillation events have on the climate in
Manu. The weather in Puerto Maldonado was hotter and possibly rainier during the
November 1997-January 1998 event, but this is not a consistent pattern, and the picture
remains foggy for much of western Amazonia (Marengo 1998).
Light
In order to measure solar radiation, a silicon pyranometer and a datalogger that records
readings every 30 minutes were established in the CICRA clearing in September 2004.
In the 257 days of data available to date, the site has received a 24-hour average of
178.8 Watts/m2. This is slightly lower than but quite similar to comparable data from
Manaus, Brazil (181), and Cristóbal, Panamá (194; Müller 1982, Cubit et al. 1988).
Seasonal variation in radiation is fairly predictable in the data to date: highest around the
equinoxes and lowest around the winter solstice.
Wind
As in many other Amazonian forests, occasional, fierce, brief windstorms play an
important role in the dynamics of Madre de Dios plant communities (Nelson et al. 1994).
Foster and Terborgh (1998) document an easterly squall that moved across Madre de
Dios in the fall of 1995, toppling thousands of trees. Ortiz (2002) describes a similar
event in the Tambopata region. On June 21, 2005, strong winds flattened a large area of
floodplain forest at the intersection of Cocha Lobo, Aguajina, and Lindero trails. The
winds were apparently associated with a very heavy rainfall event that evening, but the
CICRA wind speed meter did not record exceptionally high winds that day. Indeed, both
average wind speed and wind gust for that day do not fall in the top 100 records for the
preceding year. Taken together with the relatively local scale of damage, this suggests
that the winds involved were especially strong gusts at a particular and rather small
19
area. We do not yet know if the same event created other similar clearings elsewhere in
the area.
Satellite images and aerial photos of the Los Amigos region show at least four areas
where large patches of forests appear to have been flattened by this type of windstorm
between 1990 and 2000 (INRENA et al. 2004). All three of them are >25 ha; the largest
appear to be >150 ha. Finding four blowdowns of this size in a relatively small area
would seem to suggest that these are relatively frequent events across Madre de Dios.
However, a recent study that compared 1990 and 2000 Landsat images of a >7 million
ha area including Los Amigos watershed and Manu and Alto Purús National Parks did
not detect any other large blowdowns during that decade (INRENA et al. 2004).
In a 2000 Landsat image, three of the four blowdown patches in the Los Amigos region
have identical colors, suggesting that the disturbance there may have been
simultaneous (i.e., caused by a single storm). In the false-color Landsat these three
patches are the same color as beaches, indicating that most of their vegetation was
obliterated. The fourth patch resembles secondary forests in the image. In 2004 a visit to
this fourth patch confirmed that it has been recolonized by adult pioneer trees,
suggesting that it is several years older than the other patches (J. Olivier, personal
communication). The dominant pioneer tree in this patch, Cecropia sciadophylla
(Cecropiaceae), is the same species that dominates second growth around the CICRA
station, as well as large blowdown regeneration in northern Peru and Ecuador
(Vriesendorp et al. 2004, 2005).
Seasonality
Seasonal changes in climate, hydrology, phenology, animal behavior, and land use are
marked in Madre de Dios, but they have not yet been integrated into a single database
that permits a global view of seasonality. In the meantime, here is a brief sampling of
seasonal patterns. Large catfish migrate up the Madre de Dios River to spawn between
December and April, and largely disappear from the river for the rest of the year
(Goulding et al. 2003). Brazil nut trees put out flowers between October and January,
drop fruits in December and January, and are visited by castañeros in January and
February (Cornejo Valverde 2003). Giant otters give birth between May and June
(Schenck 1999), river turtles lay their eggs on the Río Los Amigos between July and
September (Mansilla et al. 2004), macaw visits to Tambopata clay licks peak in
September (Brightsmith 2004), and spider monkeys disperse seeds of Virola calophylla
at Cocha Cashu between September and December (Russo & Augspurger 2004). Most
tree-cutting, mining, commercial fishing, ecotourist visits and scientific research in Madre
de Dios take place during the dry season (Goulding et al. 2003).
It is not a trivial task to determine which seasonal changes drive which seasonal patterns
in plant and animal communities, since many seasonal changes occur simultaneously.
The dry season in Madre de Dios is also the season with the lowest air temperatures,
the highest river water temperatures, the lowest solar radiation levels, the thickest leaf
litter on the forest floor, the highest river water pH, the shortest days, the highest stream
conductivities, and (because it is also a time of lower rainfall in the Andes) the lowest
river levels (Goulding et al. 2003). Other seasonal patterns, like variation in air pollution,
cloud cover, wind direction and intensity, storm intensity, lightning strikes, tree falls, and
bamboo growth, are also potentially important drivers of seasonal patterns but are poorly
documented. Meanwhile, the vast majority of seasonal patterns in plant and animal
communities—changes in mycorrhizal fungi abundance, mosquito community structure,
20
disease incidence in primate populations, to name a few scattered examples—remain

undocumented. Compiling data on these seasonal patterns into a single database that
can provide a broad temporal context for research in the region, and that will allow us to
track changes in seasonality with accelerating global change, is a leading priority for
ACA.
VEGETATION AND FLORA

Regional overview and recommended bibliographic sources
The Los Amigos flora has been the subject of very intensive study and inventory by the
Andes-Amazon Botany Project since 2001, and the accumulating results will soon
provide an excellent picture of the watershed’s vegetation (J. Janovec and colleagues, in
preparation). These upcoming publications, maps, and other products should give us a
good estimate of the size of the flora, an excellent description of its composition, high-
quality maps and descriptions of the various forest types in the region, other insights into
how the plant community changes from place to place, and a great deal of other broad
and specific botanical information. In the meantime, photographs of thousands of plant
species collected at Los Amigos, as well as accompanying ecological, geographic, and
descriptive data, are available at atrium.andesamazon.org; a description of the Los
Amigos botany team’s work is available at www.andesamazon.org.
In this section I discuss a few broad topics that I consider especially important or
interesting about the region’s vegetation. Readers wanting a broad introduction to the
vegetation of the region should try Foster (1990, 1994) and Gentry (1997). For
references on more specific topics, see Table 2.
Table 2. Some recommended bibliographic sources for the vegetation of Madre de Dios, in
English and Spanish.
Key references Key references

Topic in English in Spanish
Primary succession along Kalliola et al. (1991),
Puhakka et al. (1993)
lowland rivers Terborgh & Petren (1991)
Gentry & Terborgh (1990),
Lowland floodplain forests Kvist & Nebel (2001), Kvist & Nebel (2000)
Hamilton et al. (in press)
Lowland herb and shrub
Foster (1990) Foster (1986)
communities
Bamboo-dominated forests Griscom & Ashton (2003) Silman et al. (2003)
Foster & Alban Castillo
The Pampas del Heath Foster (2002)
(1994)
Upper-elevation forests ? ?
Tree species abundance and
Pitman et al. (1999, 2001) Pitman et al. (2003, 2005)
distribution
Endemic species Pitman et al. (1999) León et al. (2007)
Secondary vegetation and
Gutte & Mueller (1989) ?
invasive species
Bush et al. (2004), Bush &
Paleobotany ?
Silman (2007)
Carbon sequestration by Naughton-Treves et al.
Chambi (2001)
forests (2004)
Fragmentation, deforestation, Phillips et al. (2006) INRENA et al. (2003, 2004)
21
and edge effects

Lawrence et al. (2005),
Alexiades (1999), Alexiades
Ethnobotany Lacaze & Alexiades (1995)
& Lacaze (1996), Phillips et
al. (1994)
History of botanical work to date at Los Amigos

(suggested 3-4 paragraphs covering dates, places, and relevant people involved in
broad floristic inventory and specific quantitative surveys; a map would be great)
The flora of Los Amigos

(suggested 2-3 pages that report the species/genus/family-level diversity recorded to
date; note especially diverse families and genera, and relative contributions to the flora
of different habit groups [e.g., trees are diverse, epiphytes aren't]; estimate prevalence of
undescribed species in the flora; hypothesize broadly to what extent the flora is similar to
those in Manu or Tambopata, and farther afield; and include a couple of floristic novelty
stories. Abundant and repeated links to Atrium. It's important, though, to keep it brief.
This isn't a summation of what we've learned but rather a quick reference for other Los
Amigos scientists who need a thumbnail sketch of plant life there.)
(Regarding phenology, that could form a brief section of its own or a couple of
paragraphs in this section.)
Vegetation of the Los Amigos watershed

(suggested 1-2-page broad-scale description of forest structure, composition, dynamics,
and heterogeneity in the watershed. Again, the trick here is not to overdo it and write
down everything we've learned but to provide a very simple description that
acknowledges its simplicity and will stand the test of time reasonably well.)
Vegetation types in the Los Amigos watershed

(suggested 2-3 paragraph description of each of the principal kinds of vegetation at Los
Amigos: mature upland forests, mature floodplain forests, aguajales, other swamps,
bamboo patches, etc. Again, the trick here is not to overdo it and write down everything
we've learned but to provide a very simple description that acknowledges its simplicity
and will stand the test of time reasonably well.)
22
Figure 5. A map of southwestern Amazonia with the extent

of bamboo-dominated upland forest, as estimated from
remote sensing imagery, shown as the cross-hatched area.
Adapted from Bianchini (2005).
Forests around CICRA

(suggested 2-3-page discussion on the flora and vegetation on the CICRA trail system.
Given that this tiny patch is the only part of the watershed that most visitors see and
collect data in, it deserves a couple of pages on its common forest types and
idiosyncrasies. For example, I want to write a few paragraphs on the stronger-than-
normal pioneer signal in forests throughout the CICRA trail system, and what may have
caused it.)
(Ideally, we'd also have sections called "Forests around CM1" and one called "Forests
around CM2". I don't know enough about those forests to contribute much, though.)
FAUNA
(under construction)
HUMAN HISTORY
Introduction
Deep History
1900-1982
We know very little about the history of the Los Amigos watershed for most of the 20th
century. In part, this is because no one has made a concerted search for mention of Los
Amigos in the hundreds of books and articles written about Madre de Dios during and
following the rubber boom. For the time being, this section relies on a few scattered
allusions I have found in geographers’ and missionaries’ accounts, and others which
anthropologist James Graham has brought to my attention.
23
One of the first published mentions of the Los Amigos River is a 1902 map of the Madre
de Dios River and tributaries by Rafael Baluarte. The map’s tracing of the Los Amigos is
cursory enough to suggest that it was drawn based on interviews rather than exploration.
While the river’s mouth is accurately mapped, its course is wildly misrepresented—
draining from east to west rather than from north to south (Figure x). Three years after
that map’s publication, the Los Amigos is described briefly in an article in the Boletín de
la Sociedad Geográfica de Lima as “a long river populated by Peruvian rubber tappers”
formed by the confluence of two smaller rivers, the Aucaio and the Busamanu
(Larrabure-Correa 1905: 92). Stiglich (1922) identifies Busamanu as the original name of
the Los Amigos, although linguistic evidence suggests that these were names given by
the rubber tappers’ guides, members of the Takana language group, rather than the
residents of the Los Amigos themselves, who more likely belonged to the Iñapari group
(T. Moore, personal communication). What Busamanu and Aucaio mean is still to be
discovered.
In a 1907 book, Stiglich confirms that the Los Amigos was one of the few tributaries of
the Madre de Dios whose forests were at that time producing for the rubber trade; rubber
tappers were unable to work in several other watersheds because of hostile indigenous
inhabitants. Indeed, the name “Amigos” is apparently a tribute to a peaceful interlude
between rubber tappers and an indigenous community that lived near the mouth of the
river. Juan M. Ontaneda (1907: 423) explains: “One of [famous rubber tapper Carlos
Fermín] Fitzcarrald’s expeditions was attacked by some [mashcos], with whom they
made peace on their return by giving them some of those cheap gifts that fascinate the
savages. As this interaction took place at the mouth of a stream, Fitzcarrald named it
‘Amigo....’”
Later in the same book, the author opens a window on earlier relations between rubber
tappers and indigenous inhabitants of the Los Amigos, which are markedly less than
friendly. He writes: “The explorers of this region had to fight with the fierce mashcos to
clean them out of the Manu and Amigos rivers, which the explorers later occupied. The
Mashcos who left the latter [apparently the Amigos] have since regrouped in the
Colorado River” (Ontaneda 1907: 449). The earliest of these “explorers” were apparently
rubber tappers led by a Peruvian named Alcibiades Torres (Stiglich 1922).
Who the indigenous inhabitants of the Amigos were and where their descendants are
living today is still not clear. Avencio Villarejo (1959) describes a settlement of 50 people
at the mouth of the Los Amigos River, inhabited by a group he calls the Manuquiaris, or
Mashcos. The first name is synonymous with the Arasairi, Sapiteri and Toyoeri of the
Amarikeri language family (J. Graham, personal communication). Ribiero and Wise
(1978) have since suggested that the Manuquiaris are most closely associated with the
Toyoeri, a group that they say has since integrated with the mestizo population of Puerto
Maldonado.
Following the battles and forced migrations of the rubber boom period, conflict between
indigenous inhabitants and mestizos has continued sporadically up to the present day
(see previous section). In a 1940 account of a trip from Puerto Maldonado to the
Colorado River, a Dominican missionary writes: “On the fourth day [of travel upriver from
the mouth of the Inambari River] we arrived at the Los Amigos River, where Señor
Troncoso and others have lived for several years. A son of his is traveling with us, and
told us that in a nearby creek the maschos had killed two of his workers and left the
bodies surrounded with arrows stuck in the ground as a sign that they would return
24
(Álvarez 1940: 177).” (The descendants of this Troncoso today own the Hotel Cabaña
Quinta in Puerto Maldonado; they believe that this story is a tall tale.)
It would appear that by the 1940s no one was tapping rubber trees any longer in the
vicinity of Los Amigos. Indeed, given that it is relatively easy to find scarred old trees in
areas of northern Amazonian Peru that were worked by rubber tappers a century ago (N.
Pitman, personal observation), the absence of scarred trees at Los Amigos is something
of a mystery. Perhaps the tapped trees died of stress or disease, or perhaps they have
yet to be found.
The Rosaurora farm plot (1974-1981)

In 1974, after their farm across the river from CICRA was flooded, the Luque family
established a farm plot at what is now CICRA. According to Juana Luque, who was still
living in the original family farm in 2006, the forest on the terrace appeared untouched
when they arrived. Seven hectares were cleared and planted with mango, avocado,
orange, and Inga trees, as well as manioc and other crops. The garden was seeded with
the African livestock grass Brachiaria, which, as Mrs. Luque noted with satisfaction
during a visit to CICRA in 2006, still grows in patches around the station. The new farm
was named Rosaurora, after a pair of the Luque family’s pet parrots, but the land was
sold shortly after to a mining company.
The mining camp (1982-1999)

Between 1982 and 1999, the site now occupied by CICRA was the center of operations
for a series of gold mining concerns. In 1982-1983 three cement buildings—what are
now CICRA’s first dormitory, dining hall, and laboratory—were constructed by a
Panamanian mining company, Texas Gulf Peru S. A. From 1983 to 1985, as many as
120 men lived in the camp, which they used as a base for explorations throughout the
region. Following the first election of Alán García as Peru’s president in 1985, the
company changed hands and operations dwindled until there were only a handful of
workers at the camp. In 1988-1990, the Peruvian company AURINSA reinitiated work at
the camp, and again the campsite was crowded with workers struggling to rebuild a
gigantic mining dredge that had been brought from Panama. This three-story dredge,
measuring some 20 x 60 m, commenced operations in 1992. Because the dredge was
mostly active upriver from CICRA, at Lago San Juan, only a few AURINSA caretakers
lived at the Los Amigos camp during 1992-2000. In 1999, amid mounting financial
problems, the mining company relocated the dredge to the Inambari River (where it sits
abandoned today) and the Los Amigos camp was stripped of its remaining assets by
frustrated employees. When ACCA purchased the property in 2000, what had been a
modern complex a few years earlier was roofless and overgrown with weeds.
Today only a few reminders of the mining camp remain: the ruins of the original winch,
standing in the yard west of the main dormitory; the cement table next to the washing
basins at the eastern end of the laboratory, which was formerly used to clean the day’s
catch of fish; the abandoned earthmoving equipment in the shed at the start of Carretera
trail; and Carretera trail itself, which the mining company built as a road for heavy
vehicles around 1992 (thus the rocks). Aerodromo trail is the continuation of this road,
built during the clearing of the landing strip that same year. The landing strip was
completed during the mining years but never used; the story has it that the only pilot to
attempt a landing blanched at the rolling terrain at the last minute and landed at Lagarto
instead.
25
The ecological impact of this large mining camp on the forests surrounding CICRA is
hard to quantify. Throughout its existence, workers at the camp fished and hunted in the
surrounding forests, because the company did not buy meat from Puerto Maldonado.
When the camp was full, providing bushmeat was a full-time job for two camp
employees. The favorite prey for hunters were deer, peccaries, tapirs, and agoutis, as
well as monkeys and large birds; the favorite fishing spot was Pañacocha. Workers cut
several large timber trees in the surrounding forest, especially tornillo (Cedrelinga
cateniformis), which was used for building, and Brazil nut (Bertholletia excelsa), which
was used for constructing mining barges. Trash was thrown over the cliff near where the
staff quarters stand today. Had 120 men lived at the camp throughout its existence, the
plant and animal communities around CICRA would have been very severely
impoverished. Thus it is a stroke of luck for biologists that the mining camp stood largely
empty for most of the 1990s.
Even so, some of those who saw the mining camp at the height of its activity today
remember it with nostalgia. In retrospect, it is easy to imagine how, from the perspective
of someone who climbed up the mining camp’s long wooden stairway, lit up with electric
lights, and stepped into a campsite thrumming with large generators, winches, and the
bustle of 120 workers, today’s CICRA is a sleepy place by comparison. Where the
cabins stand today, not long ago stood a hangar and landing pad for the helicopter that
transported the company’s engineers to and from the city. The laboratory that once
processed several kilos of gold per year is now a discolored patch in the grass near the
dining hall. The forest along today’s stairway is only a few years old; previously one
could stand at the top of the stairs and look down at the docks, or survey the river to the
south. Gone is the avocado plantation formerly around what is now the staff quarters,
gone the field of corn, manioc, and watermelon on what is now the soccer pitch, and
gone the small building that once stood at the entrance of Aerodromo trail, where the
station’s workers were entertained each month by company-sponsored visitadoras like
those described in Vargas Llosa’s novel.
The logging boom (1999-2002)
Forest structure. The impact of logging on forest structure in the Los Amigos watershed
was trivial in the period 2001-2002, so the reduction in impact is correspondingly small.
However, it is still measurable. The reasoning is as follows. Between 1 May 2001 and 30
April 2002, ACCA park guards recorded ~1.8 million board feet of mahogany and cedar
leaving the Amigos watershed. Sixty-four percent of the wood was mahogany and 36%
was cedar (ACCA 2002). Assuming 1,992 board feet/tree, and assuming that twice as
many trees were felled as were harvested (after Schulte-Herbrüggen & Rossiter 2003),
this gives a total of 1,820 large trees felled in the watershed during a one-year period:
1,156 mahogany and 664 cedar. Assuming that all of these trees were felled inside the
135,000 ha of what is now the conservation concession, this gives an impact of one tree
felled every 74 ha. Because we know that some of the wood brought out of the Los
Amigos during that time was felled outside of what is now the conservation concession,
we can round that figure to an approximate average density of one large tree felled every
100 ha.
To put this in perspective, data from a 2002 study in Cocha Cashu suggest that roughly
730 very large (>70 cm dbh) trees grow in 100 ha of unlogged forest (N. Pitman & M.
26
Silman, unpublished data). Assuming 2% mortality, an average 15 of those trees will die
of natural causes each year. By contrast, during the year of highest impact in the Los
Amigos watershed, loggers felled just one tree per 100 ha. Even if the extrapolation
exercise above underestimates impact by 100%, i.e., even if the loggers felled two trees
per 100 ha, not one -- and even if this impact occurred every year, rather than being
concentrated in one or two years -- the impact of logging remains <15% of the natural
background rates of large tree mortality. Indeed, single windstorms in this part of Peru
occasionally knock down more large trees in half an hour than were felled by loggers in
the Los Amigos watershed in 2001-2002 (Foster & Terborgh 1998, Ortiz 2002).
(Additional thoughts re this question: Juan Carlos has some data on the additional impact
of logging camps, trash, and trails. I believe these impacts are small. I can’t even find a
large logging camp I visited near CICRA in 1998, even though it was then surrounded by
very wide trails.)
Animal communities. The reduced impact of hunting is easy to estimate, by combining

ACCA’s existing data on how many people and camps were in the Los Amigos
watershed with Schulte-Herbrüggen & Rossiter’s (2003) measurements of kg bushmeat
consumed per person in logging camps on the Las Piedras River in 2002. ACCA has
already quantified these impacts in its environmental impact report to INRENA in 2002.
That document estimates that 12,884 kg of bushmeat were harvested in the Los Amigos
watershed during 2001-2002. Since the fauna and hunter preferences are similar in the
Los Amigos and Las Piedras watersheds, Schulte-Herbrüggen & Rossiter’s (2003) data
allow us to extrapolate a rough list of mammals killed during that time in the Los Amigos
watershed: 206 A. belzebuth, 179 T. pecari, 144 T. tajacu, and 107 A. seniculus. The
same sort of extrapolation can be done for birds. Compared with these numbers, zero kg
of bushmeat were harvested in the concession in 2004.
The arrival of ACCA (2000-present)
Uncontacted indigenous groups

Huertas Castillo (2002), Michael & Beier (2003)
RESEARCH AT CICRA
Research priorities
As Webb (2005) has noted, “The field of conservation biology is in a unique position: in a
world of increasing human health, wealth, and technological capacity, few other
disciplines are waging losing wars.” Faced with this gloomy tableau, ACA’s approach to
defining research priorities has been to keep things simple. Briefly, research priorities at
Los Amigos are inventory, long-term monitoring and global change, regional topics, and
public service, as described in more detail below.
• Biotic and abiotic inventories. The long-term goal is a comprehensive description

of the Los Amigos landscape, flora, and fauna, so that our managers know what
they are protecting. Outputs of inventory projects at Los Amigos range from
simple species lists to field guides, taxonomic treatments, phylogenies, on-site
synoptic collections, museum collections, and soil, vegetation, or other maps or
GIS layers. ACA favors inventory projects that focus on poorly-known groups of
27
organisms, use new technologies to maximize efficiency, share information

widely, and build links with our long-term monitoring programs. Biological
inventories to date have focused on terrestrial mammals, birds, amphibians,
reptiles, fish, various insect groups, flowering plants, lichens, mosses, and fungi
(see Table 3). Coverage is weakest, predictably, on invertebrate taxa.
28
Table 3. Biological inventories in the Los Amigos watershed, 2000-2007.
Number of species and

Taxonomic Researchers and morphospecies List, guide or publication
Group inventory dates registered to date available?
Microbial David Bass and Thomas under study in preparation

eukaryotes Richards, 2005
Ascomycete Amy Berkov, 2004-2006 6 Berkov et al. (2007)
yeasts
Fungi Romina Gazis, 2003-2005 305 Gazis (2004, 2005), Gazis
& Gómez (in preparation)
Alejandro Sánchez, 2006 under study in preparation
Lichens Jorge Carrillo, 2003 under study in preparation
Mosses Piers Majestyk, 2002 58 Majestyk & Janovec (2004)
Pteridophytes Benjamín Chambi, 2005- 176 in preparation
2006
Angiosperms John Janovec and >2,000 species list available at
colleagues, 2001-present atrium.andesamazon.org
Bivalves Cristián Ituarte, 2006 under study in preparation
Terrestrial Gabriela Cuezzo and Rina under study in preparation
gastropods Ramírez, 2006
Copepods Diana López, 2006 under study in preparation
Aquatic insects Eduardo Oyague, 2004- under study in preparation
2005
Scorpions José Antonio Ochoa, 2005 5 Ochoa & Chaparro (2005)
Spiders Mariajosé Deza, 2005-2006 under study in preparation
Flies Craig Gibbs, 2004-2005 under study in preparation
Steven Paiero & John under study in preparation
Klymko, 2006
Butterflies Phil Devries, 2003-2005 under study in preparation
Moths Pedro Centeno, 2004-2005 under study in preparation
Juan Grados, 2006 under study in preparation
Ants Various, 2004-2005 under study in preparation
Army ants in Angélico Asenjo, 2006 5 unpublished species list
the genus
Eciton
Meliponnine David Roubik, 2003 under study in preparation
and euglossine
bees
Megalopta Simon Tierney, 2006 under study in preparation
bees
Cerambycid Amy Berkov, 2004-2005 37 unpublished species list
beetles
Curculionid Amy Berkov, 2004-2005 7 unpublished species list
beetles
Erotylid beetles Dino Vanucci, 2003 28 unpublished species list
Scarabaeid Trond Larsen, 2000-2005 154 unpublished species list
beetles
Fish Michael Goulding, Carlos 292 Goulding et al. (2003),
Cañas, Hernán Ortega, Max Barthem et al. (2003)
Hidalgo, Soraya Martínez,
Vanessa Chipollini and
others, 2002-2006
Amphibians Rudolf von May and others, 82 von May (2004)
2002-2006
Reptiles Rudolf von May and others, ~50 von May (2004)
2002-2006
Birds Douglas Stotz, Tatiana 515 unpublished species list
29
Pequeño, Renán Valega,

Daniel Lebbin, and others,
2001-2006
Terrestrial and Louise Emmons, Renata 64 unpublished species list
arboreal Leite, and others, 2001-
mammals 2006
Bats Cristina López, Sandra >33 in preparation
Velazco, Adriana Bravo, and
Jhony Ríos, 2005-2006
TOTAL >3,817
• Long-term monitoring and global change. Managing the conservation concession

effectively over its 40-year lease requires an understanding of how its plant and
animal communities change over time, both naturally and because of human-
caused global change. Current monitoring projects at Los Amigos include the
collection of basic weather and river level data, weekly surveys of animal
populations along the Los Amigos River, monthly phenological, leaf-litter, and
leaf area index surveys, and multi-year surveys of permanent tree plots. Long-
term goals of the monitoring program at Los Amigos include 1) linking our
monitoring efforts to those at other stations in the region, 2) monitoring the
conservation status of globally threatened taxa, and 3) implementing
Conservation International’s standardized long-term monitoring program for
plants, butterflies, ants, birds, and primates (TEAM).
• Regional topics. ACA has a special interest in projects that study the defining
features of the southwestern Amazon landscape. These include colpas (clay
licks), bamboo-dominated forests, wetlands, friajes (winter cold fronts),
meandering rivers, economically important species like Brazil nut, mahogany,
and tropical cedar trees, endemic species, and the environmental impacts of
logging, gold mining, and ecotourism.
• Public service. Regardless of the research question, ACA favors research that
generates some public good for the community. Examples include projects that
provide training opportunities for Peruvian biologists, strengthen capacity in
Peruvian museums and universities, share research projects widely, and make a
special effort to build on prior scientific work at Los Amigos and the other well-
studied sites in the region, like Cocha Cashu, Pakitza, and the Río Tambopata
lodges. Collaborative projects with Bolivian and Brazilian researchers and
institutions are highly encouraged.
Research in Los Amigos before ACCA

The massive landslide just upriver from CICRA, identified on geological maps as Cerro
Colorado, has been studied by geologists and paleontologists since the 1980s
(Campbell & Romero Pittman 1989, Chávez et al. 1998, Campbell et al. 2000, Hovikoski
et al. 2005, Campbell et al. 2006)
In 1992, John Terborgh, Percy Núñez and Manuel Sánchez scouted forests in the Los
Amigos watershed as part of a large-scale inventory of tree communities in Madre de
Dios. Terborgh and colleagues had completed several tree inventories in the Manu
watershed and were inspecting tree communities in other watersheds in Madre de
Dios—an undertaking that took them to the Alto Madre de Dios, the Colorado, the
30
Pariamanu, the Las Piedras, the Tahuamanu and the Acre rivers, as well as the Los
Amigos. In 1993, Núñez, Sánchez and Viviana Horna returned to Los Amigos to
establish two 2-ha tree plots in upland forest. In 1998, Núñez, Sánchez and Nigel Pitman
visited the two plots as part of Pitman’s thesis work. One plot, now located in upland
forest near CICRA between Castañal and Segundo Mirador trails, was recensused by
Rolando Díaz in 2005. The other plot, located in upland forest near CM3, was relocated
and recensused by Carlos Lazo and the Andes-Amazon Botany Project team in early
2007. Data from these plots have appeared in several articles and theses (Terborgh &
Andresen 1998, Pitman 2000, Pitman et al. 2001, Condit et al. 2001, Pitman et al. 2002,
Terborgh et al. 2002, ter Steege et al. 2004, Masse 2005, Latimer et al. 2006, Malhi et
al. 2006).
In 2000, Wake Forest master’s student Emilio Ancaya sampled woody vegetation the
large palm swamp northwest of CICRA (Ancaya 2002). Miles Silman, Mark Bush, and
Luis Imunda accompanied Ancaya as part of a project to sample swamp vegetation and
pollen in lake sediments in the Manu and Madre de Dios watersheds.
Stern/Cornejo trip
FZS surveys
The first six years (2000-2006)

Research at CICRA began shortly after the purchase of the former mining camp by
ACCA in 2000. The first researchers to work at Los Amigos under the auspices of ACCA
were Trond Larsen and Alejandro Lopera, who surveyed dung beetles at CICRA in April
and May 2000. Larsen recalled that: “The laboratory building was... overgrown and did
not have any roof.... Our dung beetle laboratory consisted of a makeshift table under a
tree in front of the dining hall (see Figure 6).”
Figure 6. The CICRA laboratory building in April 2000 (left) and September 2006 (right). Photos:
Trond Larsen and Nigel Pitman.
Relatively few researchers visited the station in 2000-2003 as its infrastructure, staff, and
management were being put into place (Figure 7). In each of those four years, the
number of person-days of research averaged <2,000; in other words, fewer than five
researchers and assistants were present at the station per day, on average, during that
period. Notable projects included the start of the long-term botanical inventory by Dr.
John Janovec, in 2001, a variety of rapid biological inventories (plants, birds, mammals)
for the elaboration of the proposal conservation concession in 2001, and an intensive
31
inventory of fish communities and aquatic habitats around the station in 2001-2002, by
Dr. Michael Goulding and colleagues (Goulding et al. 2003, Barthem et al. 2003).
Research traffic accelerated sharply in early 2004 as grantees from the new ACA grants
program began arriving at the station and a large radio-tracking project led by Dr.
George Powell got underway. During 2004 the average number of researchers and
assistants rose to 18; by 2005 this number had risen to 31.
Figure 7. Growth in research activity at Los Amigos, 2000-2006.
Since 2003, ACA’s grants program to support field research at Los Amigos has made
111 awards—an average of more than one grant every two weeks (Table 4). ACA has a
special interest in providing support to Peruvian and Latin American scientists.
Consequently, 60% of grants to date have been awarded to Peruvian researchers, and
>15% of the remainder have been awarded to researchers from other Latin American
countries. ACA offers three kinds of support: 1) grants to Peruvian undergraduates
working on their thesis project, 2) grants to Peruvian and foreign graduate students
working on their thesis project, and 3) grants to Peruvian and foreign established
researchers starting the first year of a multi-year project at Los Amigos.
Table 4. Grants awarded by ACA’s program to support field research at Los Amigos, 2003-2007.
Figures in parenthesis are the number of awards given to Peruvian grantees.
Undergraduate Graduate Seed All

Year grants grants grants grants
2003 8 (8) 3 (2) 2 (0) 13 (10)
2004 9 (9) 11 (3) 10 (4) 30 (16)
2005 8 (8) 8 (3) 10 (6) 26 (17)
32
2006 4 (4) 7 (2) 10 (5) 21 (11)

2007 7 (7) 7 (1) 7 (4) 21 (12)
TOTALS 36 (36) 36 (11) 39 (19) 111 (66)
33
ACKNOWLEDGMENTS
I am indebted to Edgar Collado, Carlos Cañas, Pascual Flores, Viviana Horna, Trond
Larsen, Juana Luque, Miles Silman, the ACCA promotores, and many others who have
shared information and observations. This document should be cited as: Pitman, N. C.
A. 2010. An overview of the Los Amigos watershed, Madre de Dios, southeastern Peru.
February 2010 version of an unpublished report available from the author at
npitman@amazonconservation.org.
34
ANNOTATED BIBLIOGRAPHY
The following combines a list of references cited in this article and a list of all known
publications related to Los Amigos. The latter are marked by stars.
*Abrill, C., S. Alvarado, A. Luque, R. Martinez and L. Portal. 2004. Riqueza de invertebrados
acuáticos asociados a brácteas de Heliconia stricta (Musaceae): Una aproximación a la
teoría de biogeografía de islas. Pages 35-37 in Servat, G. (ed.), unpublished
compendium of student projects from the 2004 field course "Ecología tropical y diseño
experimental" at the Centro de Investigación y Capacitación del Río Los Amigos,
sponsored by the Asociación para la Conservación de la Cuenca Amazónica (ACCA).
*Abrill, C., N. Hidalgo and S. Zambrano. 2004. Riqueza de aves en bosque de tierra firme e
inundable. Pages 45-49 in Servat, G. (ed.), unpublished compendium of student projects
from the 2004 field course "Ecología tropical y diseño experimental" at the Centro de
Investigación y Capacitación del Río Los Amigos, sponsored by the Asociación para la
Conservación de la Cuenca Amazónica (ACCA).
*ACCA. 2000. Diagnóstico situacional de las comunidades de Boca Amigo, Centro Poblado
Menor de San Juan Grande y Cinco Islas. Unpublished report of the Asociación para la
Conservación de la Cuenca Amazónica.
*ACCA. 2001a. Plan de negocios para el Centro de Capacitación Los Amigos. Unpublished
report of the Asociación para la Conservación de la Cuenca Amazónica.
*ACCA. 2001b. Propuesta técnica para una concesión de conservación en la cuenca del río de
Los Amigos, departamento de Madre de Dios. Unpublished report presented to INRENA
by the Asociación para la Conservación de la Cuenca Amazónica (ACCA). 42 pages.
*ACCA. 2002. Declaración de impacto ambiental, Concesión para Conservación en la Cuenca
del Río Los Amigos. Unpublished report presented to the Instituto Nacional de Recursos
Naturales (INRENA) by the Asociación para la Conservación de la Cuenca Amazónica
(ACCA). 30 pages.
*ACCA. 2003. Informe anual 2002-2003, concesión para conservación en la cuenca del Río Los
Amigos. Unpublished report of the Asociación para la Conservación de la Cuenca
Amazónica (ACCA). Lima.
*Achicahuala, J., V. Cano, C. González, J. Vargas, L. Zenteno and D. Cadena. 2003. Cúanto
tienes, cuánto vales: Disponibilidad de recursos y la defensa en las plantas. Pages 96-99
in Servat, G., M. Rodríguez, D. Cadena, K. Balta, and C. García-Robledo (eds.),
unpublished compendium of student projects from OTS course "Ecología de ecosistemas
amazónicos 2003-13," sponsored by the Universidad Nacional de la Amazonía Peruana,
Centro Amazónico de Educación Ambiental e Investigación & the Organization for
Tropical Studies.
*Alberdi, M. T., J. L. Prado and R. Salas. 2004. The Pleistocene Gomphotheriidae (Proboscidea)
from Peru. Neues Jahrbuch für Geologie und Palaontologie-Abhandlungen 231(3): 423-
452. ABSTRACT: Morphological characters of South American Gomphothere remains
from Peru are described and discussed in this work. We assign the remains from
Ayusbamba (Cuzco) to Cuvieronius hyodon and the remains from La Huaca (Piura),
Quipan (Canta) and Serpentin of Pasamayo to Stegomastodon waringi. It is not possible
to distinguish remains from Rio Madre de Dios, referred to the late Miocene of Amazonia,
from other Stegomastodon waringi specimens from late Pleistocene of Peru based on
dental morphology. Consequently, further data are needed to confirm the earliest
appearance of Gomphotheriidae in Peru. We also describe Gomphothere migration
routes from the North and examine their palaeobiogeographical implications.
*Alfaro, A., S. Alvarado, N. Andrade, E. Chulla and A. Luque. 2004. Relación entre la riqueza y la
abundancia de insectos y la cobertura en un bosque de succesión ribereña. Pages 57-61
in Servat, G. (ed.), unpublished compendium of student projects from the 2004 field
course "Ecología tropical y diseño experimental" at the Centro de Investigación y
Capacitación del Río Los Amigos, sponsored by the Asociación para la Conservación de
la Cuenca Amazónica (ACCA).
35
*Alonso Castillo, F. 2001. Evaluación de impacto ambiental de la concesión para la conservación

de cuenca del Río Los Amigos. Unpublished report of the Asociación para la
Conservación de la Cuenca Amazónica. Lima. 137 pages.
*Alvarez, A., K. Balta, L. Campos, C. del Aguila, T. P. Feria, L. Nolivos and T. Erwin. 2002. ¿Qué
hay más allá del orificio?: Riqueza y abundancia de los insectos acuáticos presentes en
los internodos del bambú Guadua weberbaueri. Pages 88-91 in Servat, G., S. Correa, A.
Bravo, and D. Cadena (eds.), unpublished compendium of student projects from OTS
course "Ecología de ecosistemas amazónicos 2002-13," sponsored by the Universidad
Nacional de la Amazonía Peruana and the Organization for Tropical Studies.
*Alvarez, M. and L. D. Avila. 2002. Evaluación de la hipótesis de concentración de recursos en
Duguetia flagellaris (Annonaceae) en el bosque de terraza del Centro “Los Amigos”.
Pages 134-138 in Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.), unpublished
compendium of student projects from OTS course "Ecología de ecosistemas amazónicos
2002-13," sponsored by the Universidad Nacional de la Amazonía Peruana and the
Organization for Tropical Studies.
*Amable, R. 2004. Avistamiento y monitoreo de nidos de guacamayo cabeza azul Propyrrhura
couloni, CICRA, Perú. Unpublished report of the WWF AREAS project for the Amazon
Conservation Association. 3 pages.
*Antoine, P.-O., P. Baby, S. Brusset, W. Hermoza and M. Roddaz. 2003. Informe de campo:
Geología de la Concesión Los Amigos: Misión de reconocimiento (setiembre del 2003).
Unpublished report for the Asociación para la Conservación de la Cuenca Amazónica
(ACCA). 26 pages.
*Aragón, I. 2003. Comportamiento de Saguinus imperator en Rio Los Amigos: Reporte de avance
del 05/II al 15/IV del 2003. Unpublished report for the Asociación para la Conservación
de la Cuenca Amazónica (ACCA). 9 pages.
*Araujo, A., J. Curo, G. Namen, C. Sánchez, A. M. Vasco-P., M. Zalamea, G. W. Fernandes and
A. Tauro. 2004. Tricomas como mecanismo de protección contra herbivoría. Pages 87-
90 in Servat, G., D. Cadena, K. Balta, and C. García-Robledo (eds.), unpublished
compendium of student projects from the OTS course "Ecología de ecosistemas
Tropical Studies.
*Arnaiz Perales, J. Year unknown. Ordenamiento del área de recursos naturales del Bosque de
los Niños (BoNi) de la comunidad de Boca Amigos. Unpublished report of the Asociación
para la Niñez y su Ambiente. Puerto Maldonado.
*Artavia, G., A. M. Vasco-P., R. A. Saldaña, M. Gómez, A. Araujo and E. Fischer. 2004. Efecto de
la carga alar en el potencial de dispersión en semillas de Banisteriopsis sp.
(Malpighiaceae). Pages 84-86 in Servat, G., D. Cadena, K. Balta, and C. García-Robledo
(eds.), unpublished compendium of student projects from the OTS course "Ecología de
ecosistemas amazónicos 2004-13," sponsored by the Universidad Nacional de la
Amazonía Peruana, Centro Amazónico de Educación Ambiental e Investigación & the
*Aschero, V., L. Zenteno, S. Ríos, C. González, J. Achicahuala, J. Vargas and T. Erwin. 2003. La
misma obra, diferentes actores: Estructura y composición de las comunidades de
insectos acuáticos en los internudos del bambú (Guadua weberbaueri). Pages 87-90 in
Servat, G., M. Rodríguez, D. Cadena, K. Balta, and C. García-Robledo (eds.),
Tropical Studies.
Avencio Villarejo O.S.A. 1959. La selva y el hombre. Editorial Ausonia, Lima.
*Avila, L. D., L. Y. Cazoria, V. Méndez, C. L. Quan, S. Vejarano, R. Dirzo, O. Vargas and F.
Cornejo. 2002. Inversión en defensa contra herbivoría en plantas de crecimiento rápido y
lento. Pages 92-100 in Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.),
amazónicos 2002-13," sponsored by the Universidad Nacional de la Amazonía Peruana
36
and the Organization for Tropical Studies.

*Azá, J. P. 1927. Hidrografía del departamento de Madre de Dios. Misiones Dominicanas del
Perú 9(41): 147-151.
*Barthem, R., M. Goulding, B. Forsberg, C. Cañas and H. Ortega. 2003. Aquatic ecology of the
Río Madre de Dios: Scientific bases for Andes-Amazon headwaters conservation.
Asociación para la Conservación de la Cuenca Amazónica/Amazon Conservation
Association, Lima. 117 pages.
*Barthem, R., M. Goulding, B. Forsberg, C. Cañas and H. Ortega. 2003. Ecología acuática del
Río Madre de Dios: Bases científicas para la conservación de cabeceras andino-
amazónicas. Asociación para la Conservación de la Cuenca Amazónica/Amazon
Conservation Association, Lima. 117 pages.
*Beltrán, H. 2001. Informe de prospección botánica en la cuenca del río Los Amigos.
(ACCA).
*Berkov, A. 2003. From Los Amigos: Wood-boring beetles associated with the castaña family
(Lecythidaceae). Unpublished report for the Asociación para la Conservación de la
Cuenca Amazónica (ACCA).
Bianchini, M. C. Florestas dominadas por bambu (gênero Guadua) no sudoeste da Amazônia:
Extensão, comportamento espectral e associação com o relevo. 2005. Master’s thesis.
Instituto Nacional de Pesquisas da Amazônia & Universidade do Amazonas, Manaus. 75
pages.
*Blundo, C., A. Tauro and J. Carilla. 2004. Estructura y variaciones morfológicas de dos especies
de plantas del dosel en bosques de terraza alta y terraza baja en Madre de Dios. Pages
117-120 in Servat, G., D. Cadena, K. Balta, and C. García-Robledo (eds.), unpublished
Tropical Studies.
*Botero, R., J. L. Flores, C. González, A. Horta, D. C. Ramos and M. Aizen. 2002. Efecto de la
posición de la flor sobre la dilución del néctar después de un chubasco. Pages 77-82 in
Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.), unpublished compendium of
student projects from OTS course "Ecología de ecosistemas amazónicos 2002-13,"
sponsored by the Universidad Nacional de la Amazonía Peruana and the Organization
for Tropical Studies.
*Botero, R. and A. Horta. 2002. Hojas rojas: ¿Una estrategia de escape a la herbivoria? Pages
101-105 in Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.), unpublished
2002-13," sponsored by the Universidad Nacional de la Amazonía Peruana and the
Brightsmith, D. J. 2004. Effects of weather on avian geophagy in Tambopata, Peru. Wilson
Bulletin 116: 134-145.
Brightsmith, D. J. and R. A. Munoz-Najar. 2004. Avian geophagy and soil characteristics in southeastern
Peru. Biotropica 36(4): 534-543.
*Bruner, A. 2001. Preliminary findings on the economics of small scale logging in the Rio Amigos.
Unpublished report for the Amazon Conservation Association (ACA). 2 pages.
*Campbell, K. E., C. D. Frailey and L. R. Romero-Pittman. 2000. The late Miocene Gomphothere
Amahuacatherium peruvium (Proboscidea: Gomphotheriidae) from Amazonian Peru:
Implications for the great American faunal interchange. Pages INGEMMET, Boletin No
23, Serie D.
*Campos, L. and C. del Aguila. 2002. Asociación entre la ocurrencia de patógenos y la herbivoría
en la lámina foliar de Banara guianensis (Flacourtiaceae). Pages 150-153 in Servat, G.,
S. Correa, A. Bravo, and D. Cadena (eds.), unpublished compendium of student projects
from OTS course "Ecología de ecosistemas amazónicos 2002-13," sponsored by the
Universidad Nacional de la Amazonía Peruana and the Organization for Tropical Studies.
*Cañas, C. 2001. Un primer acercamiento a la ictiofauna alrededor del centro río Los Amigos.
37
(ACCA).
*Carilla, J., R. Giudice, A. M. Torres, K. Eckhardt, K. Dexter and G. W. Fernandes. 2004. Efectos
de la filogenia y el área foliar sobre el nivel de ataque y la efectividad en la respuesta
ante insectos agalladores en plantas. Pages 70-75 in Servat, G., D. Cadena, K. Balta,
and C. García-Robledo (eds.), unpublished compendium of student projects from the
OTS course "Ecología de ecosistemas amazónicos 2004-13," sponsored by the
Universidad Nacional de la Amazonía Peruana, Centro Amazónico de Educación
Ambiental e Investigación & the Organization for Tropical Studies.
*Cazoría, M. Y. and V. Méndez Alvarez. 2002. Mimercofilia y herbivoría en Triplaris americana y
Triplaris poeppigiana (Polygonaceae) en la reserva Los Amigos, Perú. Pages 127-133 in
Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.), unpublished compendium of
*Chave, J., H. C. Muller-Landau, R. Condit, N. Pitman, J. Terborgh, S. P. Hubbell and E. G.
Leigh. 2002. Beta-diversity in tropical forests: Response. Science 297(5586).
*Chávez V., A., G. Salas A., J. Cuadros P. and E. Gutiérrez S. (eds.). 1998. Boletín No. 106,
Serie A. Carta Geológica Nacional: Geología de los cuadrángulos de Fitzcarrald y Río
Los Amigos (Hojas 25-u y 25-v). Instituto Geológico Minero y Metalurgico, Lima.
*Chocce, M., J. P. Janovec and E. A. Christenson. 2004. A synopsis of the genus Otostylis
(Orchidaceae: Maxillarieae subtribe Zygopetalinae) with a new record from southern
Peru. Sida 21(2): 841-852. ABSTRACT: The genus Otostylis (Orchidaceae) is one of the
lesser known genera in the Zygopetalum alliance, which has had a problematic
taxonomic history. Otostylis consists of four species distributed in South America and
Trinidad. The purpose of this paper is to provide a synopsis of the genus, with
descriptions of the four species and keys for their identification. Otostylis paludosa is
resurrected from previous synonomy under O. lepida and reported as a new record for
the genus in Peru, occurring as a dominant herb in bog wetlands of the Department of
Madre de Dios.
*Chocce Peña, M. A. 2003. Estudio de las orquídeas de tres bosques de la cuenca del Río Los
Amigos (Dpto. Madre de Dios, Perú). Unpublished report for the Asociación para la
Conservación de la Cuenca Amazónica. 6 pages.
*Condit, R., N. Pitman, E. G. Leigh, J. Chave, J. Terborgh, R. B. Foster, P. Nunez, S. Aguilar, R.
Valencia, G. Villa, H. C. Muller-Landau, E. Losos and S. P. Hubbell. 2002. Beta-diversity
in tropical forest trees. Science 295(5555): 666-669. ABSTRACT: The high alpha-
diversity of tropical forests has been amply documented, but beta-diversity -- how species
composition changes with distance -- has seldom been studied. We present quantitative
estimates of beta-diversity for tropical trees by comparing species composition of plots in
lowland terra firme forest in Panama, Ecuador, and Peru. We compare observations with
predictions derived from a neutral model in which habitat is uniform and only dispersal
and speciation influence species turnover. We find that beta-diversity is higher in Panama
than in western Amazonia and that patterns in both areas are inconsistent with the
neutral model. In Panama, habitat variation appears to increase species turnover relative
to Amazonia, where unexpectedly tow turnover over great distances suggests that
population densities of some species are bounded by as yet unidentified processes. At
intermediate scales in both regions, observations can be matched by theory, suggesting
that dispersal limitation, with speciation, influences species turnover.
*Conger, J. 2001. Organic gardening at Los Amigos. Unpublished report for the Asociación para
la Conservación de la Cuenca Amazónica (ACCA).
Cornejo Valverde, F. 2003. Historia natural de la castaña (Bertholletia excelsa Humb. & Bonpl.) y
propuestas para su manejo. Asociación para la Conservación de la Cuenca Amazónica,
Puerto Maldonado. 52 pages.
*Cortez, C. and D. C. Ramos. 2002. Diferencias en los niveles de herbivoria entre sexos en una
planta dioica (Cecropia sciadophylla). Pages 106-110 in Servat, G., S. Correa, A. Bravo,
and D. Cadena (eds.), unpublished compendium of student projects from OTS course
"Ecología de ecosistemas amazónicos 2002-13," sponsored by the Universidad Nacional
38
de la Amazonía Peruana and the Organization for Tropical Studies.

*Costantino, M. and A. Troya. 2003. Efecto de los cambios en la intensidad de luz sobre la
abundancia y morfología foliar de Vanilla pompona (Orchidaceae). Pages 127-129 in
Tropical Studies.
Cubit, J. D., R. C. Thompson, H. M. Caffey, and D. M. Windsor. 1988. Hydrographic and
meteorological studies of a Caribbean fringing reef at Punta Galeta, Panamá: Hourly and
daily variations for 1977-1985. Smithsonian Contributions to the Marine Sciences 32: 1-
220.
*Cuenca, V., J. Chahuara and L. Alvarez. 2004. Factores que determinan la riqueza de insectos
en claros y bosque. Pages 53-56 in Servat, G. (ed.), unpublished compendium of student
projects from the 2004 field course "Ecología tropical y diseño experimental" at the
Centro de Investigación y Capacitación del Río Los Amigos, sponsored by the
Asociación para la Conservación de la Cuenca Amazónica (ACCA).
*Cuenca, V., N. Hidalgo, E. Valderrama and S. Zambrano. 2004. Depredación denso
dependiente en semillas de Iriartea deltoidea (Arecaceae). Pages 27-30 in Servat, G.
(ed.), unpublished compendium of student projects from the 2004 field course "Ecología
tropical y diseño experimental" at the Centro de Investigación y Capacitación del Río Los
Amigos, sponsored by the Asociación para la Conservación de la Cuenca Amazónica
(ACCA).
*Curo, A. and A. Fernández. 2003. Corre gota corre: Función del acúmen en el drenaje de agua
en las plantas. Pages 108-110 in Servat, G., M. Rodríguez, D. Cadena, K. Balta, and C.
García-Robledo (eds.), unpublished compendium of student projects from OTS course
de la Amazonía Peruana, Centro Amazónico de Educación Ambiental e Investigación &
the Organization for Tropical Studies.
*Curo, J. L. and A. J. Araujo. 2004. Distribución temporal de ortopteros asociados a Paspalum
sp. (Poaceae). Pages 124-128 in Servat, G., D. Cadena, K. Balta, and C. García-
Robledo (eds.), unpublished compendium of student projects from the OTS course
*Curo, J. L., M. Espinoza, M. Rivera, A. Tauro, I. Yánez, M. Zalamea and T. Erwin. 2004. Efecto
de tiempo de establecimiento sobre la composición de comunidades de artrópodos
asociados a Tessaria integrifolia. Pages 80-83 in Servat, G., D. Cadena, K. Balta, and C.
García-Robledo (eds.), unpublished compendium of student projects from the OTS
Nacional de la Amazonía Peruana, Centro Amazónico de Educación Ambiental e
Investigación & the Organization for Tropical Studies.
Davenport, L. 2003. La problemática de las aguas: Los lobos de río y los hábitats acuáticos de la
Zona Reservada del Alto Purús. Pages 125-136 in R. Leite Pitman, N. Pitman, and P.
Álvarez (eds.), Alto Purús: Biodiversidad, conservación y manejo. Center for Tropical
Conservation, Lima.
*de la Torre Cuba, A. 2003. Informe final de campo: Diversidad genética entre poblaciones de
Cedrela odorata: Implicancias para la conservación. Unpublished report for the
Asociación para la Conservación de la Cuenca Amazónica (ACCA). 4 pages.
*de la Torre Cuba, A. R. M. 2005. Diversidad genética entre poblaciones de Cedrela odorata:
Implicancias para la conservación. Master's thesis. Escuela de Post-Grado,
Especialización en Conservación de Recursos Forestales, Universidad Nacional Agraria
La Molina, Lima. 70 pages.
*Delgado, C., A. Costantino, N. Estrada, C. Fagua, J. Machuca and T. Erwin. 2003. Especie
clave para el establecimiento de fauna acuática en los internudos del bambú:
¿Escarabajo o carpintero? Pages 100-102 in Servat, G., M. Rodríguez, D. Cadena, K.
39
Balta, and C. García-Robledo (eds.), unpublished compendium of student projects from

*Dexter, K., A. M. Vasco-P. and M. Zalamea. 2004. ¿Son los troncos en descomposición islas de
recursos para hongos en el contexto de la teoría de biogeografía de islas? Pages 105-
108 in Servat, G., D. Cadena, K. Balta, and C. García-Robledo (eds.), unpublished
Tropical Studies.
*Dubois, S. 2004. Etude des modalités de croissance et de dispersion des populations de
bambou (Guadua weberbaueri) entrant en compte lors de la colonisation de zones
forestières tropicales perturbées par l'activité humaine dans le sud-est péruvien.
Unpublished report of ECOFIT (Ecosystèmes et paléoécosystèmes des Forêts
Intertropicales). 74 pages.
*Eckhardt, K., C. Rodríguez, D. Rodríguez, M. Rivera, M. Espinoza, A. M. Torres, E. Fischer and
J. Carilla. 2004. Abundancia y riqueza de ácaros asociados a flores de seis especies de
plantas visitadas por colibríes. Pages 91-94 in Servat, G., D. Cadena, K. Balta, and C.
García-Robledo (eds.), unpublished compendium of student projects from the OTS
*Ellison, K. 2003. Renting: The conservation concessions approach. Conservation Biology in
Practice 4(4): 20-29.
*Emmons, L. H. 2001. Mammals of Los Amigos. Unpublished report for the Asociación para la
Conservación de la Cuenca Amazónica.
*Emmons, L. H. and M. A. Dubois. 2003. Leaf-area index change across river-beach
successional transects in south-eastern Peru. Journal of Tropical Ecology 19: 473-477.
Emmons, L. H. and N. M. Stark. 1979. Elemental composition of a natural mineral lick in
Amazonia. Biotropica 11(4): 311-313.
Erwin, T. L. 1985. Tambopata Reserved Zone, Madre de Dios, Peru: History and description of
the reserve. Revista Peruana de Entomología 27: 1-8.
*Estrada, N., A. Curo, M. Costantino, J. Díaz, J. Rodríguez and J. Janovec. 2003. Efecto de la
profundidad del agua y la luz en la distribución de plántulas de aguaje (Mauritia
flexuosa). Pages 75-78 in Servat, G., M. Rodríguez, D. Cadena, K. Balta, and C. García-
Robledo (eds.), unpublished compendium of student projects from OTS course "Ecología
de ecosistemas amazónicos 2003-13," sponsored by the Universidad Nacional de la
*Etienne, R. S., A. M. Latimer, J. A. Silander, Jr. and R. M. Cowling. 2006. Comment on "Neutral
ecology theory reveals isolation and rapid speciation in a biodiversity hot spot". Science
311: 610b.
*Fagua, C., C. Cárdenas, J. Machuca, M. Rodríguez, Y. Paiz and D. Cadena. 2003. Ensamblaje
de comunidades de artrópodos en hojas secas de Cecropia membranacea:
¿Determinismo o azar? Pages 83-86 in Servat, G., M. Rodríguez, D. Cadena, K. Balta,
and C. García-Robledo (eds.), unpublished compendium of student projects from OTS
*Fagua, C., C. Delgado and A. Gil. 2003. Establecimiento de epífitas en las raíces de Socratea
exorrhiza e Iriartea deltoidea. Pages 130-134 in Servat, G., M. Rodríguez, D. Cadena, K.
40
*Feria, T. P. and K. Balta. 2002. Efecto del volumen de agua sobre la riqueza de invertebrados
presente en los internodos del bambú Guadua weberbaueri. Pages 117-122 in Servat,
G., S. Correa, A. Bravo, and D. Cadena (eds.), unpublished compendium of student
projects from OTS course "Ecología de ecosistemas amazónicos 2002-13," sponsored
by the Universidad Nacional de la Amazonía Peruana and the Organization for Tropical
Studies.
*Fernández, M. Year unknown. Informe final de la prospección ambiental del río de Los Amigos.
(ACCA).
*Ferreyra Vela, F. R. 2004. Conservación del lobo de río (Pteronura brasiliensis) en Perú,
Capítulo II: Censo de población de nutria gigante (Pteronura brasiliensis) en las cuencas
de los ríos Amigos y bajo Madre de Dios. Unpublished report of the Proyecto Lobo de
Río of the Frankfurt Zoological Society for the Amazon Conservation Association. 35
pages.
*Flores, C. 2004. Effect of light, soil fertility and soil moisture in early growth of lowland tropical
cedars (Cedrela spp.) in southeast Peru. Unpublished report for the International
Foundation of Science.
*Flores, J. L. and O. González. 2002. Efectividad del camuflaje en saltamontes. Pages 145-149
in Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.), unpublished compendium of
*Forsyth, A., M. Ríos and M. Romo. 2002. Plan de manejo: Concesión para conservación en la
cuenca del río Los Amigos, provincias de Manu y Tambopata, Departamento de Madre
de Dios. Unpublished report presented to the Instituto Nacional de los Recursos
Naturales (INRENA) by the Asociación para la Conservación de la Cuenca Amazónica
(ACCA). 82 pages.
Foster, M. S. and J. Terborgh. 1998. Impact of a rare storm event on an Amazonian forest.
Biotropica 30(3): 470-474.
Foster, R. B. 1990. Long-term change in the successional forest community of the Rio manu
floodplain. Pages 565-572 in A. H. Gentry (ed.), Four Neotropical rainforests. Yale
University Press, New Haven, USA.
*Foster, R. B. 2001. Some description of the Río Los Amigos, Madre de Dios, Peru. Unpublished
report for the Asociación para la Conservación de la Cuenca Amazónica (ACCA). 2
pages.
*Foster, R. B., H. Betz and H. Beltrán. 2002. Arboles y arbustos del Centro Río Amigos. An
unpublished laminated field guide produced by Environmental and Conservation
Programs of the Field Museum. 30 + 600 color photos pages.
*Gazis Oliva, R. 2004. Evaluación preliminar de la micoflora localizada en los alrededores del
Centro de Investigación "Río Los Amigos," Manu. Tesis de licenciatura. Facultad de
Ciencias Biológicas, Universidad Ricardo Palma, Lima.
*Gil, A., V. Cano, C. Delgado, V. Fernández, A. Montero, A. Troya and R. Dirzo. 2003. Morfología
foliar como mecanismo de escape a herbívoros en Iriartea deltoidea. Pages 79-82 in
Tropical Studies.
*Giudice, R., C. Blundo, J. Carilla, G. Artavia, F. Soley, T. Erwin and C. Sánchez. 2004.
Composición de comunidades de artrópodos en tres tipos de vegetación de sucesión
ribereña. Pages 95-99 in Servat, G., D. Cadena, K. Balta, and C. García-Robledo (eds.),
unpublished compendium of student projects from the OTS course "Ecología de
ecosistemas amazónicos 2004-13," sponsored by the Universidad Nacional de la
*Giudice, R., G. Namen and R. A. Saldaña. 2004. Segregación espacial de murciélagos y
41
características morfológicas en sotobosques cerrados y abiertos. Pages 109-112 in

Servat, G., D. Cadena, K. Balta, and C. García-Robledo (eds.), unpublished compendium
of student projects from the OTS course "Ecología de ecosistemas amazónicos 2004-
13," sponsored by the Universidad Nacional de la Amazonía Peruana, Centro Amazónico
de Educación Ambiental e Investigación & the Organization for Tropical Studies.
*Gonzáles, T., M. Chocce, K. Siu Ting, J. Chahuara and L. Alvarez. 2004. Relación entre la
riqueza de insectos y la biomasa de la vegetación en un bosque de succesión ribereña
del río Madre de Dios, Perú. Pages 31-34 in Servat, G. (ed.), unpublished compendium
of student projects from the 2004 field course "Ecología tropical y diseño experimental" at
the Centro de Investigación y Capacitación del Río Los Amigos, sponsored by the
*Gonzáles, T., R. Martínez, K. Siu Ting and N. Medina. 2004. Uso de hábitats por juveniles y
adultos de Paleosuchus palpebrosus (Alligatoridae). Pages 39-44 in Servat, G. (ed.),
unpublished compendium of student projects from the 2004 field course "Ecología
tropical y diseño experimental" at the Centro de Investigación y Capacitación del Río Los
Amigos, sponsored by the Asociación para la Conservación de la Cuenca Amazónica
(ACCA).
*González, C. and F. Ferreyra. 2002. Efecto de la distancia entre plantas sobre el robo de néctar
en Besleria sp. (Gesneriaceae). Pages 154-160 in Servat, G., S. Correa, A. Bravo, and
D. Cadena (eds.), unpublished compendium of student projects from OTS course
de la Amazonía Peruana and the Organization for Tropical Studies.
*Goulding, M., C. Cañas, R. Barthem, B. Forsberg and H. Ortega. 2003. Amazon headwaters:
Rivers, wildlife, and conservation in southeastern Peru. Grafica Biblos S.A., Lima. 117
pages.
*Goulding, M., C. Cañas, R. Barthem, B. Forsberg and H. Ortega. 2003. Las fuentes del
Amazonas: Ríos, vida y conservación del sureste peruano. Asociación para la
Conservación de la Cuenca Amazónica/Amazon Conservation Association, Lima.
*Gutiérrez, N. 2002. Diversidad florística en aguajales Mauritia flexuosa L. f. (Arecaceae).
Unpublished report of unknown provenance.
*Hardner, J. and R. Rice. 2002. Rethinking green consumerism. Scientific American 286: 89-95.
*Hovikoski, J., M. Rasanen, M. Gingras, M. Roddaz, S. Brusset, W. Hermoza and L. R. Pittman.
2005. Miocene semidiurnal tidal rhythmites in Madre de dios, Peru. Geology 33(3): 177-
180. ABSTRACT: New data from upper Miocene deposits in the Madre de Dios region,
southern Peru, allow the delineation of tidal regime for the first time in western Amazonia
and provide strong evidence of elevated tidal range and brackish-water influence. The
results point out the insufficiency of the current depositional models and support the
earlier hypothesis that western Amazonia was also connected to the Paranan Sea during
the late Miocene. In this paper we present sedimentological, ichnological, and statistical
(Fourier transformation) data from two selected outcrops containing rhythmite
successions from an area that is traditionally considered as continental. The sediments
are interpreted to represent tide-dominated, inner-middle estuarine deposits. The cyclic
rhythmites display semidiurnal cyclicity. The results are significant because (1) they
contradict recent interpretations of the area's paleoenvironmental history; (2) the
evidence for tidal processes is persuasive; and (3) the delineated tidal regime and range
provide a unique insight into the depositional dynamics of a system having many
important paleogeographic implications.
*Huamaní, G. 2000. Cuenca del río Los Amigos: Mapa base y fisiografía "base de datos
georeferenciada". Unpublished report of unknown provenance.
Huertas Castillo, B. 2002. Los pueblos indígenas en aislamiento: Su lucha por la sobrevivencia y
la libertad. Tarea Gráfica Educativa, Lima.
*INRENA, SZF and CDC-UNALM. 2004. Hacia un sistema de monitoreo ambiental remoto
estandarizado para el SINANPE: Estudio de caso 2003: Parque Nacional Manu y Zona
Reservada del Alto Purús. Unpublished report of the Instituto Nacional de Recursos
Naturales (INRENA), Sociedad Zoológica de Francfort (SZF), and the Centro de Datos
para la Conservación (CDC-UNALM). Lima. 133 pages.
42
Kalliola, R., and M. Puhakka. 1993. Geografía de la selva baja peruana. Pages 9-21 in R.
Kalliola, M. Puhakka, and W. Danjoy (eds.), Amazonía peruana: Vegetación húmeda
tropical en el llano subandino. PAUT and ONERN, Jyväskylä, Finland.
Kalliola, R., M. Puhakka and W. Danjoy (eds.). 1993. Amazonía peruana: Vegetación húmeda
tropical en el llano subandino. Proyecto Amazonia of the Universidad de Turku (PAUT),
and Oficina Nacional de Evaluación de Recursos Naturales (ONERN), Jyväskylä. 261
pages.
*Kirkby, C. 2001. The distribution, abundance, clump characteristics and techniques for managing
Guadua cf. angustifolia, Bambuseae, a potential non-wood forest product, in Madre de
Dios, Peru. Master's thesis. The University of York, York, UK.
*Kirkby, C. and B. Griscom. 2001. Bamboo research and development strategy for the Los
Amigos Conservation Concession and associated areas. Unpublished report for the
Asociación para la Conservación de la Cuenca Amazónica. 29 pages.
*Kometter, R. F., M. Martinez, A. G. Blundell, R. E. Gullison, M. K. Steininger and R. E. Rice.
2004. Impacts of unsustainable mahogany logging in Bolivia and Peru. Ecology and
Society 9(1): 12. ABSTRACT: Although bigleaf mahogany [Swietenia macrophylla King
(Meliaceae)] is the premier timber species of Latin America, its exploitation is
unsustainable because of a pattern of local depletion and shifting supply. We surveyed
experts on the status of mahogany in Bolivia and Peru, the world's past and present
largest exporters. Bolivia no longer has commercially viable mahogany (trees > 60 cm
diameter at breast height) across 79% of its range. In Peru, mahogany's range has
shrunk by 50%, and, within a decade, a further 28% will be logged out. Approximately
15% of the mahogany range in these two countries is protected, but low densities and
illegal logging mean that this overestimates the extent of mahogany under protection.
The international community can support mahogany conservation by funding park
management and by encouraging independent verification of the legality of mahogany in
trade. Our findings demonstrate that a systematic expert survey can generate reliable
and cost-effective information on the status of widespread species of concern and help to
inform appropriate management policy.
*Küchmeister, H. 2000. Pasos para un plan de manejo sostenible de la cuenca del río Los
Amigos. Unpublished report for the Asociación para la Conservación de la Cuenca
Amazónica.
*La Rosa, D. 2002. Patrones de uso estacional de las colpas por vertebrados mayores en el
Centro de Investigación Río Los Amigos, Madre de Dios. Unpublished report for the
*La Rosa, D. 2003. Patrones de uso de collpas por mamíferos en el Río Los Amigos, Madre de
Dios. Unpublished report for the Asociación para la Conservación de la Cuenca
Amazónica. 11 pages.
*La Rosa, D., L. H. Emmons, M. Trolle and V. Pacheco (2003). Uso de colpas por el tapir de
tierras bajas Tapirus terrestris (Perissodactyla: Tapiridae) en el Río Los Amigos, Madre
de Dios. XII Reunión Científica. Universidad Nacional Mayor de San Marcos, April 2003,
Lima.
Larsen, M. C., and A. Simon. 1993. A rainfall intensity-duration threshold for landslides in a
humid-tropical environment, Puerto Rico. Geografiska Annaler 75A: 13-23.
*Larsen, T. 2000. Los Amigos dung beetle report. Unpublished report for the Asociación para la
Conservación de la Cuenca Amazónica. 6 pages.
*Latimer, A. M., J. A. Silander, Jr. and R. M. Cowling. 2005. Neutral ecological theory reveals
isolation and rapid speciation in a biodiversity hot spot. Science 309: 1722-1725.
ABSTRACT: South Africa’s Mediterranean-climate fynbos shrubland is a hot spot of
species diversity, but its diversity patterns contrast strongly with other high-diversity
areas, including the Amazon rain forest. With its extremely high levels of endemism and
species turnover, fynbos is made up of dissimilar local communities that are species-rich
but relatively poor in rare species. Using neutral ecological theory, we show that the
relative species-abundance distributions in fynbos can be explained by migration rates
that are two orders of magnitude lower than they are in tropical rain forests. Speciation
rates, which are indexed by the ‘‘biodiversity parameter’’ Q, are estimated to be higher
43
than they are in any previously examined plant system.

*Lawrence, J. M. 2003. Preliminary report on the natural history of brown titi monkeys (Callicebus
brunneus) at the Los Amigos Research Station, Madre de Dios, Peru. American Journal
of Physical Anthropology: 136-136.
*Lebbin, D. 2003. 2003 field season progress report: Spatial and structural aspects of bamboo
that promote specialization among Amazonian birds. Unpublished report for the
*Leite Pitman, R. and G. Powell. 2004. Reporte anual del estudio de áreas utilizadas por algunas
especies claves de mamíferos y aves grandes en la cuenca del río Los Amigos.
Unpublished report of the WWF AREAS project for the Amazon Conservation
Association. 36 pages.
Leopold, A. [1924] 1991. The river of the mother of God and other essays. University of
Wisconsin Press, Madison, USA.
Linna, A. 1993. Factores que contribuyen a las caracteristicas del sedimento superficial en la
selva baja de la Amazonia peruana. Pages 87-97 in R. Kalliola, M. Puhakka, and W.
Danjoy (eds.), Amazonia peruana: Vegetación húmeda tropical en el llano subandino.
PAUT and ONERN, Jyväskylä, Finland.
*Lipa Mercado, D. 2005. Inventario de cinco especies de palmeras en el Bosque de los Niños
(BONI) en la comunidad de Boca Amigo en la Provincia del Manu. Prácticas Pre-
profesionales report for the Facultad de Ciencias Forestales y Medio Ambiente of the
Universidad Nacional San Antonio Abad del Cusco. Puerto Maldonado. 23 pages.
*Machuca, J., C. Cárdenas and J. Díaz. 2003. Estructura de la población de Mauritia flexuosa en
un gradiente de sucesión. Pages 135-138 in Servat, G., M. Rodríguez, D. Cadena, K.
*Majestyk, P. and J. P. Janovec. 2004. Contributions to the moss flora of the Amazonian lowlands
of Madre de Dios, Peru. Sida 21(2): 1231-1238. ABSTRACT: A recent collection of
mosses from the Department of Madre de Dios in the lowlands of southeastern Peru
contains 58 different species. Fifty-four taxa are new to the district and four species are
reported new to Peru. Previous collections from the district are listed which brings the
total species known from the area to 74. This is the first major study of mosses from
Madre de Dios, Peru, and it provides a foundation for continued work and production of a
field guide for the region.
*Mansilla, A. M., E. Quispe and D. F. Ttito. 2004. Uso del recurso taricaya (Podocnemis unifilis)
en la Concesión para Conservación Río Los Amigos. Unpublished report for the Amazon
*Mansilla, A. M. and D. Ttito. 2005. Determinación de la densidad relativa de la taricaya
(Podocnemis unifilis) en cochas del Río Los Amigos. Unpublished report for the Amazon
*Mansilla U., A. M. 2004. Estudio, monitoreo y manejo de la taricaya (Podocnemis unifilis) en la
Concesión para Conservación Río Los Amigos, Madre de Dios, Perú. Unpublished report
for the Amazon Conservation Association. 19 pages.
Marengo, J., A. Cornejo, P. Satyamurty, and C. Nobre. 1997. Cold surges in tropical and
extratropical America: The strong event in June 1994. Monthly Weather Review 125 (11):
2759-86.
Marengo, J.A. 1998. Climatología de la zona de Iquitos. Pages 35-57 in R. Kalliola and S. Flores-
Paitán (eds.), Geoecología y desarrollo Amazónico: Estudio integrado en la zona de
Iquitos, Perú. Annales Universitatis Turkuensis Ser A II 144, Turku, Finland.
*Masse, D. 2005. The effects of distance and geomorphology on the floristic composition of
lowland tropical tree communities. Master's thesis. Department of Biology, Wake Forest
University, Winston-Salem, USA. ABSTRACT: A 53-tree plot network from southeastern
Peru was used to evaluate the effects of niche-based and dispersal-based processes on
the floristic composition of communities from four environmentally distinct but
geographically close geomorphological units: floodplain, successional forest, swamp, and
44
terra firme. The tree plots were arranged in floristic space using non-metric
multidimensional scaling ordination, and the effects of distance and geomorphology on
floristic similarity were evaluated using multiple linear regressions. The effect of habitat
was further explored by comparing observed habitat restriction with a null expectation.
The geomorphologies were floristically distinct, and species were more habitat-restricted
than expected by chance. There was a significant effect of distance on floristic similarity,
most of it at very short scales (< 5 km). Geomorphology explained most of the variation at
an intermediate scale (< 50 km) and distance explained most of the variation at a larger
scale (up to 300 km). Distance explained little (~5%) of the across-habitat similarity, while
it explained more than 30% within habitats. While distance affected floristic composition,
its effects were overridden by geomorphology. The large scale distance effect may be
due to the spatial variation in fluvial dynamics. The short scale distance effect probably
reflects dispersal and mass effects. Geomorphology may locally be a valid proxy for
environmental variables but seems to fail to summarize environmental variation at large
scales.
Mazer, S. J. 1997. Floristic composition, soil quality, litter accumulation, and decomposition in
terra firme and floodplain habitats near Pakitza, Peru. Pages 89-126 in D. E. Wilson and
A. Sandoval (eds.), Manu: The biodiversity of southeastern Peru. Smithsonian Institution
and Editorial Horizonte, Lima.
*Medina, N., N. Andrade, A. Alfaro and E. Chulla. 2004. Estructura de la comunidad de insectos
en tres tipos de fitotelmata. Pages 23-26 in Servat, G. (ed.), unpublished compendium of
student projects from the 2004 field course "Ecología tropical y diseño experimental" at
Michael, L. D., and C. Beier. 2003. Poblaciones indígenas en aislamiento voluntario en la región
del Alto Purús. Pages 149-162 in R. Leite Pitman, N. Pitman, and P. Álvarez (eds.), Alto
Purús: Biodiversidad, conservación y manejo. Center for Tropical Conservation, Lima.
*Montero, A., N. Estrada and Y. Paiz. 2003. ¿Qué factores afectan la riqueza de epífitas en
jardines de hormigas? Pages 118-121 in Servat, G., M. Rodríguez, D. Cadena, K. Balta,
and C. García-Robledo (eds.), unpublished compendium of student projects from OTS
*Mori, S. 2001. Key to the Lecythidaceae of Los Amigos. Unpublished report for the Asociación
para la Conservación de la Cuenca Amazónica (ACCA). 2 pages.
Müller, M. J. 1982. Selected climatic data for a global set of standard stations for vegetation
science. W. Junk, The Hague, The Netherlands.
Napravnik Pesce, M. (ed.) 2004. Guía interpretativa para Tambopata Research Center y Posada
Amazonas. 11ra Edición 1994-2004. Rainforest Expeditions S. A. C. and INNOVA Arquitectura
Integral S. R. L., Lima. 332 pages.
*Neill, A. K. Year unknown. Los Amigos jungle cucumbers: A study of five sympatric Gurania
species. Unpublished report for the Asociación para la Conservación de la Cuenca
Amazónica.
Nelson, B. W., V. Kapos, J. B. Adams, W. J. Oliveira, O. P. G. Braun, and I. L. do Amaral. 1994.
Forest disturbance by large blowdowns in the Brazilian Amazon. Ecology 75 (3): 853-
858.
*Nolivos, L. and S. Enriquez. 2002. Remoción de semillas por murciélagos y aves en plantas del
género Piper. Pages 123-126 in Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.),
*Ochoa, J. A. and J. C. Chaparro. 2005. Scorpiones de la cuenca del río Madre de Dios, Perú.
Unpublished report for the Asociación para la Conservación de la Cuenca Amazónica. 19
pages.
*Olivier, J. 2003. Distribución, historia y futuro de la invasión del bambú en la cuenca del río Los
Amigos en relación a la variación climática: Implicaciones para la tala sostenible a largo
45
plazo. Unpublished report for the Asociación para la Conservación de la Cuenca

Amazónica. 6 pages.
*Olivier, J., M. Dubois and L. Emmons (2003). A spatio-temporal analysis of flowering in Guadua
bamboo. British Ecological Society and Association for Tropical Biology Special
Symposium, University of Aberdeen, 7-10 July 2003.
ONERN. 1965. Inventario y evaluación del potencial económico social de la zona de Kosñipata-
Alto Madre de Dios-Manu. ONERN, Lima. 138 pages.
ONERN. 1972. Inventario, evaluacíon e integracíon de los recursos naturales de la zona de los
ríos Inambari y Madre de Dios. Oficina Nacional de Evaluacíon de Recursos Naturales
(ONERN), Lima. 296 pages.
ONERN. 1977. Inventario, evaluacion y integracion de los recursos naturales de la zona Iberia–
Iñapari. Oficina Nacional de Evaluacíon de Recursos Naturales (ONERN), Lima. 207
pages.
ONERN. 1982. Inventario y evaluación semidetallada de los recursos de suelos y forestales de la
zona de Atalaya. ONERN, LIMA. 177 pages.
Osher, L. J. and S. W. Buol. 1998. Relationship of soil properties to parent material and
landscape position in eastern Madre de Dios, Peru. Geoderma 83(1-2): 143-166.
*Oyague Passuni, E. 2004. El concepto de continuidad del río y su aplicabilidad en algunos
sistemas hidrográficos Neotropicales: Informe preliminar, primera salida. Unpublished
report for the Asociación para la Conservación de la Cuenca Amazónica (ACCA). 6
pages.
*Paiz, Y., V. Aschero, C. Cárdenas, A. Gil, A. Montero, M. Rodríguez and J. Janovec. 2003.
Composición de helechos en un gradiente abiótico en el aguajal. Pages 91-95 in Servat,
G., M. Rodríguez, D. Cadena, K. Balta, and C. García-Robledo (eds.), unpublished
2003-13," sponsored by the Universidad Nacional de la Amazonía Peruana, Centro
Amazónico de Educación Ambiental e Investigación & the Organization for Tropical
Studies.
Paredes Arce, G., S. Kauffman, and R. Kalliola. 1998. Suelos aluviales recientes de la zona
Iquitos-Nauta. Pages 231-252 in R. Kalliola and S. Flores-Paitán (eds.), Geoecología y
desarrollo Amazónico: Estudio integrado en la zona de Iquitos, Perú. Annales
Universitatis Turkuensis Ser A II 144, Turku, Finland.
Phillips, O. L., P. N. Vargas, A. L. Monteagudo, A. P. Cruz, M. E. C. Zans, W. G. Sanchez, M. Yli-
Halla and S. Rose. 2003. Habitat association among Amazonian tree species: a
landscape-scale approach. Journal of Ecology 91(5): 757-775.
*Pitman, N., J. Terborgh, M. P. Núñez V. and M. Valenzuela. 2003. Los árboles de la cuenca del
río Alto Purús. Pages 53-62 in R. Leite Pitman, N. Pitman and P. Álvarez (eds.), Alto
Purús: Biodiversidad, conservación y manejo. Center for Tropical Conservation, Lima.
*Pitman, N. C. A. 2000. A large-scale inventory of two Amazonian tree communities. Ph.D.
dissertation. Department of Botany, Duke University, Durham, USA. ABSTRACT: Two
tree communities at the western margin of the Amazon basin were inventoried via
networks of small plots scattered over several thousand square kilometers of forest.
Yasuní National Park, Ecuador, is a moist, aseasonal, hyperdiverse lowland forest near
the equator. Manu National Park, Peru, ~1,400 km and eleven degrees to the south of
Yasuní, is a moist, seasonal, moderately diverse lowland forest. The aims of the research
were to document patterns of distribution, abundance, and diversity among Amazonian
tree species at a variety of different spatial scales, and to investigate explanations for the
patterns. The communities were surprisingly similar in composition and structure. Most
species encountered in the inventories are believed to grow throughout western
Amazonia, and a large proportion of them occur in both plot networks. Species common
at one site tend to be common at the other, and the palm Iriartea deltoidea dominates
both forests at identical densities. Most species in these forests occur preferentially in
one forest type, but probably fewer than 15-26% of species are restricted to a single
forest type. The great majority of species are rare at all spatial scales, but at both sites a
small proportion of common taxa account for >50% of individual trees across the terra
firme landscape. Common species tend to be large-statured trees, and are especially
46
prevalent in the families Arecaceae, Myristicaceae, Moraceae, and Violaceae. The

Ecuadorean forest is more diverse than the Peruvian forest at all taxonomic levels and all
spatial scales. It also has a higher stem density, a larger proportion of smaller-statured
species, a larger proportion of rare species, and higher-than-expected increases in the
diversity of certain families. I argue that a large component of species composition and
structure in these forests is homogeneous, and demonstrate that the tree communities in
unvisited plots are largely predictable without any reference to local environmental
conditions. However, I suggest that local processes related to the higher rainfall and
higher stem density in Ecuador are more likely causing its higher diversity than large-
scale or historical influences. Many of the observed patterns remain unexplained.
*Pitman, N. C. A., M. del C. Loyola Azáldegui, K. Salas, G. Vigo T. and D. A. Lutz. (in press, July
2006). Written accounts of an Amazonian landscape over the last 450 years.
Conservation Biology.
*Pitman, N. C. A., J. Terborgh, P. Núñez and M. R. Silman. 2001. Especies arbóreas comunes de
la parte baja de Madre de Dios, Perú. Pages 46-52 in L. O. Rodríguez (ed.), El Manu y
otras experiencias de investigación y manejo de bosques neotropicales. Proyecto
Aprovechamiento y Manejo Sostenible de la Reserva de Biosfera y Parque Nacional del
Manu (PRO-MANU), Lima.
*Pitman, N. C. A., J. W. Terborgh, M. R. Silman, P. Nunez, D. A. Neill, C. E. Ceron, W. A.
Palacios and M. Aulestia. 2001. Dominance and distribution of tree species in upper
Amazonian terra firme forests. Ecology 82(8): 2101-2117. ABSTRACT: Amazonian
forests are the largest and most diverse in the tropics, and much of the mystery
surrounding their ecology can be traced to attempts to understand them through tiny local
inventories. In this paper we bring together a large number of such inventories scattered
across immense areas of western Amazonia in order to address simple questions about
the distribution and abundance of tropical tree species in lowland terra firme forests
there. The goal is to describe patterns of commonness and rarity at local (1 ha),
landscape (~10(4) km-2), and regional (>10(6) km-2) scales, and to fuse the results into a
more complete picture of how tropical tree communities are structured. We present
estimates of landscape-scale densities for ~1400 taxa, based on data from tree plots
scattered over large tracts of terra firme forest in eastern Ecuador and southeastern
Peru. A database of morphological, ecological, and other traits of >1000 of these species
compiled from the taxonomic literature is then used to explore how species that are
common in the inventories differ from species that are rare. Although most species show
landscape-scale densities of <1 individual/ha, most trees in both forests belong to a small
set of ubiquitous common species. These common species combine high frequency with
high local abundance, forming predictable oligarchies that dominate several thousand
square kilometers of forest at each site. The common species comprising these
oligarchies are a nonrandom subset of the two floras. At both sites a disproportionate
number of common species are concentrated in the families Arecaceae, Moraceae,
Myristicaceae, and Violaceae, and large-statured tree species are more likely to be
common than small ones. Nearly a third of the 150 most common tree species in the
Ecuadorean forest are also found among the 150 most common tree species in the
Peruvian forest. For the 254 tree species shared by the two data sets, abundance in
Ecuador is positively and significantly correlated with abundance ~1400 km away in Peru.
These findings challenge popular depictions of Amazonian vegetation as a small-scale
mosaic of unpredictable composition and structure. Instead, they provide additional
evidence that tropical tree communities are not qualitatively different from their temperate
counterparts, where a few common species concentrated in a few higher taxa can
dominate immense areas of forest. We hypothesize that most Amazonian forests are
dominated at large scales by oligarchies similar in nature to the ones observed in
Ecuador and Peru, and we argue that the patterns are more indicative of regulation of
relative abundances by ecological factors than of nonequilibrium chance-based
dynamics. The paper concludes with a discussion of the practical applications of
predictable oligarchies over large areas of unexplored forest.
*Pitman, N. C. A., J. W. Terborgh, M. R. Silman, P. Nunez, D. A. Neill, C. E. Ceron, W. A.
47
Palacios and M. Aulestia. 2002. A comparison of tree species diversity in two upper
Amazonian forests. Ecology 83(11): 3210-3224. ABSTRACT: We inventoried two
Amazonian tree communities separated by similar to1400 km of continuous lowland
tropical forest, in an effort to understand why one was more diverse than the other.
Yasuni National Park, near the equator in eastern Ecuador, has one of the most diverse
tree communities in the world. Manu National Park, at 12degrees S in Peru's Madre de
Dios region, is only moderately diverse by upper Amazonian standards. Following the
field inventories, a database of. morphological, ecological, and other traits was compiled
from the taxonomic literature for 1039 species from the plots. Our goals were (1), to
describe how terra firme tree communities at the two sites differed in composition,
diversity, and structure; (2) to characterize the "extra" species responsible for the higher
diversity at Yasuni; and (3) to assess, in the light of those observations, some
explanations for why forests near the equator are so diverse. Yasuni has similar to 1.4
times as many. tree species as Mann at all three spatial scales we examined: local (1
ha), landscape (<10000 km(2)), and regional (<100000 km(2)). Yasuni samples contain
more families and genera, more individual trees per unit area, and a larger proportion of
small trees. Tree species at Yasuni have smaller stature, larger leaves, larger seeds, and
smaller geographic and altitudinal ranges than those at Manu, and disproportionate
increases in species diversity are observed within the Myrtaceae, Lauraceae,
Melastomataceae, and several other families. Community structures were strikingly
similar, with the same species (Iriartea deltoidea, a palm) dominating both sites at
identical densities. Common species at Yasuni occur at the same densities as equally
ranked species at Manu, but there are substantially more very rare species at Yasuni.
The poorer tree flora is not a nested subset of the richer tree flora, though a majority of
species in each inventory do occur at the other site. Several models that offer
explanations for geographic variation in tropical tree species diversity are assessed in
light of these data. Most do a poor job of accounting for. the patterns revealed by the
inventories. We speculate that the most important factor in producing the higher diversity
in Yasuni is its rainier, aseasonal climate, and we discuss. two specific rainfall-related
mechanisms that appear to be supported by the data: (1) year-round water availability
allowing more species to persist in the understory at Yasuni and (2) a newly described
"mixing effect" related to the higher stem density there.
*Puhakka, M. and R. Kalliola. 1995. Floodplain vegetation mosaics in western Amazonia.
Biogeographica 71(1): 1-14.
*Quan, C. L. and M. Vidaurre. 2002. Composición de las comunidades de epífitas en jardines de
hormigas. Pages 139-144 in Servat, G., S. Correa, A. Bravo, and D. Cadena (eds.),
*Rado Quispe, R. 2003. Influencia de la disponibilidad de luz bajo el dosel, en el crecimiento de
regeneración natural de Bertholletia excelsa H.B.K. en tres tipos de bosques en el Centro
de Investigación del Río Los Amigos, Madre de Dios, Perú. Thesis for the degree of
Ingeniero Forestal. Universidad Nacional San Antonio Abad del Cusco, Puerto
Maldonado. 59 pages.
*Rado Quispe, R. Year unknown. Inventario de Cedrelinga catenaeformis Ducke en el Centro de
Investigación del Río Los Amigos, Madre de Dios. Prácticas Pre-profesionales report for
the Facultad de Ciencias Forestales y Medio Ambiente of the Universidad Nacional San
Antonio Abad del Cusco. Puerto Maldonado. 22 pages.
Räsänen, M. 1993. La geohistoria y geología de la Amazonia peruana. Pages 43-67 in R.
Kalliola, M. Puhakka, and W. Danjoy (eds.), Amazonia Peruana: Vegetación húmeda
tropical en el llano subandino. PAUT and ONERN, Jyväskylä, Finland.
Riley, M. P. 1994. Soil chemical changes accompanying a primary riparian succession in Manu
National Park, Madre de Dios, Peru. Master's thesis, Duke University.
*Ríos, S., J. Díaz, V. Fernández, J. Rodríguez, A. Troya and G. Servat. 2003. Riqueza y
abundancia de insectos asociados a Heliconia stricta en relación a la posición de la
bráctea. Pages 103-107 in Servat, G., M. Rodríguez, D. Cadena, K. Balta, and C. García-
48
Robledo (eds.), unpublished compendium of student projects from OTS course "Ecología
*Rodríguez, C., K. Eckhardt and M. Rivera. 2004. Abundancia y riqueza de ácaros presentes en
colibríes y en flores visitadas por éstos. Pages 121-123 in Servat, G., D. Cadena, K.
the OTS course "Ecología de ecosistemas amazónicos 2004-13," sponsored by the
*Rodríguez, D., F. Soley, C. Blundo, G. Namen, C. Sánchez, C. Rodríguez and P. Stevenson.
2004. Dispersión de semillas de Onychopetalum sp. (Annonaceae) en un bosque tropical
de la Amazonía peruana. Pages 76-79 in Servat, G., D. Cadena, K. Balta, and C. García-
*Rodríguez, J. and V. Aschero. 2003. ¿Me ves o no me ves?: Comportamiento críptico
antidepredación en ortópteros. Pages 111-113 in Servat, G., M. Rodríguez, D. Cadena,
K. Balta, and C. García-Robledo (eds.), unpublished compendium of student projects
*Rodríguez, M. and V. Cano. 2003. Efecto de la densidad y presencia de tricomas sobre la
herbivoría en Melastomataceae. Pages 114-117 in Servat, G., M. Rodríguez, D. Cadena,
K. Balta, and C. García-Robledo (eds.), unpublished compendium of student projects
*Rojas, J. and S. Vejarano. 2002. Relación entre la tasa de producción de néctar y la frecuencia
de visitas de murciélagos a flores de Bauhinia tarapotensis. Pages 111-116 in Servat, G.,
S. Correa, A. Bravo, and D. Cadena (eds.), unpublished compendium of student projects
Universidad Nacional de la Amazonía Peruana and the Organization for Tropical Studies.
*Rojas, J., M. Vidaurre, C. Cortez, O. Gonzáles, S. Enriquez, F. Ferreyra and E. Fischer. 2002. El
cambio de una forma esférica hacia una forma plana: Un posible camino evolutivo para
las diásporas anemocóricas? Pages 83-87 in Servat, G., S. Correa, A. Bravo, and D.
Cadena (eds.), unpublished compendium of student projects from OTS course "Ecología
Amazonía Peruana and the Organization for Tropical Studies.
Russo, S. E. and C. K. Augspurger. 2004. Aggregated seed dispersal by spider monkeys limits
recruitment to clumped patterns in Virola calophylla. Ecology Letters 7: xxx-xxx.
Schenck, C. 1999. Lobo de río (Pteronura brasiliensis): Presencia, uso del hábitat y protección en
el Perú. GTZ & INRENA. Lima. 176 pages.
*Salcedo Villamar, E. 2002. Evaluación en transectos de Bertholletia excelsa H.B.K. en tres
zonas de la cuenca del Río Los Amigos. Prácticas Pre-profesionales report for the
Facultad de Ciencias Forestales y Medio Ambiente of the Universidad Nacional San
Antonio Abad del Cusco. Puerto Maldonado.
*Servat, G. 2003. Ecología tropical y diseño experimental. Unpublished compendium of student
projects from a 2003 field course at the Centro de Investigación y Capacitación del Río
Los Amigos, sponsored by the Asociación para la Conservación de la Cuenca
Amazónica (ACCA). 86 pages.
*Servat, G. 2004. Ecología tropical y diseño experimental. Unpublished compendium of student
projects from a 2004 field course at the Centro de Investigación y Capacitación del Río
Los Amigos, sponsored by the Asociación para la Conservación de la Cuenca
Amazónica (ACCA). 72 pages.
49
*Servat, G., D. Cadena, K. Balta and C. García-Robledo. 2004. Ecología de ecosistemas

amazónicos OET 2004-13. Unpublished compendium of student projects from OTS
course 2004-13, sponsored by the Universidad Nacional de la Amazonía Peruana,
Tropical Studies. 138 pages.
*Servat, G., S. Correa, A. Bravo and D. Cadena. 2002. Ecología de ecosistemas amazónicos
2002-13. Unpublished compendium of student projects from OTS course 2002-13,
for Tropical Studies. 170 pages.
*Servat, G., M. Rodríguez, D. Cadena, K. Balta and C. García-Robledo. 2003. Ecología de
ecosistemas amazónicos 2003-13. Unpublished compendium of student projects from
OTS course 2003-13, sponsored by the Universidad Nacional de la Amazonía Peruana,
Tropical Studies. 164 pages.
*Soley, F., G. Artavia and C. Sánchez. 2004. Cambios en la composición y riqueza de epífitas en
jardines de hormigas de distinta edad y tamaño. Pages 113-116 in Servat, G., D.
Cadena, K. Balta, and C. García-Robledo (eds.), unpublished compendium of student
projects from the OTS course "Ecología de ecosistemas amazónicos 2004-13,"
sponsored by the Universidad Nacional de la Amazonía Peruana, Centro Amazónico de
Educación Ambiental e Investigación & the Organization for Tropical Studies.
*Stern, M. J. 1985. Bamboo-dominated forest in southwest Amazon: Are they shrinking or
expanding into surrounding heterogenous tropical lowland forest? Unpublished
manuscript.
*Stern, M. J. and F. Cornejo Valverde. 1998. Floristic structure and composition of bamboo-
dominated forests in southeastern Peru. Unpublished report prepared for WWF-US,
Botanical Gaps in Latin America and the Caribbean Program.
*Swarner, M. 2001. Resumen de 12 días de observaciones de mamíferos en el Centro del Río
Los Amigos. Unpublished report for the Asociación para la Conservación de la Cuenca
Amazónica.
*Swarner, M. Year unknown. Colpas of the Los Amigos area. Unpublished report for the
Asociación para la Conservación de la Cuenca Amazónica.
*Swift, B., S. Bass, V. Sanjinés, V. Theulen, M. Milano, M. L. Nunes, V. Maldonado, A. Cortés, C.
M. Chacón, V. Arias, M. Tobar, M. Gutierrez and P. Solano. 2003. Legal tools and
incentives for private lands conservation in Latin America: Building models for success.
Environmental Law Institute, Washington, DC, USA. 217 pages.
*ter Steege, H., N. Pitman, D. Sabatier, H. Castellanos, P. Van der Hout, D. C. Daly, M. Silveira,
O. Phillips, R. Vasquez, T. Van Andel, J. Duivenvoorden, A. A. De Oliveira, R. Ek, R.
Lilwah, R. Thomas, J. Van Essen, C. Baider, P. Maas, S. Mori, J. Terborgh, P. N. Vargas,
H. Mogollon and W. Morawetz. 2003. A spatial model of tree alpha-diversity and tree
density for the Amazon. Biodiversity and Conservation 12(11): 2255-2277. ABSTRACT:
Large-scale patterns of Amazonian biodiversity have until now been obscured by a
sparse and scattered inventory record. Here we present the first comprehensive spatial
model of tree alpha-diversity and tree density in Amazonian rainforests, based on the
largest-yet compilation of forest inventories and bolstered by a spatial interpolation
technique that allows us to estimate diversity and density in areas that have never been
inventoried. These data were then compared to continent-wide patterns of rainfall
seasonality. We find that dry season length, while only weakly correlated with average
tree alpha-diversity, is a strong predictor of tree density and of maximum tree alpha-
diversity. The most diverse forests for any given DSL are concentrated in a narrow
latitudinal band just south of the equator, while the least diverse forests for any given
DSL are found in the Guayana Shield and Amazonian Bolivia. Denser forests are more
diverse than sparser forests, even when we used a measure of diversity that corrects for
sample size. We propose that rainfall seasonality regulates tree alpha-diversity and tree
density by affecting shade tolerance and subsequently the number of different functional
types of trees that can persist in an area.
50
Terborgh, J. 1983. Five New World primates: A study in comparative ecology. Princeton
University Press, Princeton, USA.
Terborgh, J. 1990. An overview of research at Cocha Cashu Biological Station. Pages 48-59 in A.
H. Gentry (ed.), Four Neotropical rainforests. Yale University Press, New Haven, USA.
Terborgh, J., C. H. Janson, and M. Brecht. 1986. Cocha Cashu: Su vegetación, clima y recursos.
Pages 1-18 in M. A. Ríos (ed.), Reporte Manu. Centro de Datos para la Conservación,
Universidad Nacional Agraria La Molina, Lima.
*Terborgh, J. and E. Andresen. 1998. The composition of Amazonian forests: Patterns at local
and regional scales. Journal of Tropical Ecology 14: 645-664. ABSTRACT: An analysis
was conducted of floristic patterns contained in 48 1-ha tree plots distributed at 29 sites in
seven neotropical countries, with a primary emphasis on the Amazonian region. Analyses
were made with family level data, using detrended correspondence analysis and
multidimensional scaling to generate two-dimensional ordinations. Dissimilarity values for
all pairs of plots were then used to compare forest composition at both local (flooded vs
unflooded forests) and regional scales (e.g., western vs central vs eastern Amazonia).
The predominate family of trees in a large majority of Amazonian and Guianan forests (by
number of stems) is either Palmae or Leguminosae (sensu latu), followed by Moraceae
and Euphorbiaceae. The forests of western Amazonia are particularly rich in palms,
Moraceae, and Myristicaceae, whereas those of eastern Amazonia and the Guianas are
rich in Lecythidaceae and Chrysobalanaceae. Dissimilarity between sites increases with
distance for both flooded and unflooded forests. The tree communities of flooded and
unflooded forests within a region tended to resemble one another more closely than
forests of either type resembled the homologous forests of the adjoining regions. Within
Amazonia the edaphic properties of each region and its geological history are tightly
interrelated. it is therefore difficult to distinguish between evolutionary and ecological
interpretations of the results.
*Terborgh, J., N. Pitman, M. Silman, H. Schichter and P. Núñez V. 2002. Maintenance of tree
diversity in tropical forests. Pages 1-17 in D. J. Levey, W. Silva and M. Galetti (eds.),
Seed dispersal and frugivory: Ecology, evolution and conservation. CAB International,
Oxford University Press, Oxford, UK.
*Torres, A. M., D. Rodríguez and M. Espinoza. 2004. Selección de entrenudos de bambú por
Rhinastus latisternus (Curculionidae). Pages 129-133 in Servat, G., D. Cadena, K. Balta,
*Trolle, M. 2003. Camera trapping in the Rio Los Amigos region, Madre de Dios, Amazonia, SE
Peru. Unpublished report for the Asociación para la Conservación Amazónica.
*Trolle, M. 2004. Flashed in the forest. BBC Wildlife Magazine 22: 30-37.
*Trolle, M. and L. H. Emmons. 2004. A record of a dwarf brocket from lowland Madre de Dios,
Peru. Deer Specialist Group News 19: 2-5. ABSTRACT: The South American dwarf
brockets (Mazama spp.) are known from few museum specimens, and their geographic
ranges and population status are poorly known (Wemmer 1998; IUCN 2002). We report
here on a dwarf brocket photographed by camera trap in the Madre de Dios region of SE
Peru. We believe that this is the first published record of a dwarf brocket in the lowland
Amazonian rainforest of Peru and, possibly, the first unambiguous record of dwarf
brocket from all of the Amazonian lowlands. It seems to be a new species record for Peru
and comprises either a major range extension of a known species or, possibly, an
undescribed taxon.
*Trolle, M. and M. Kéry. 2004. Camera-trap study of ocelot and other secretive mammals in the
Peruvian Amazon. Unpublished report for the Amazon Conservation Association. 5
pages.
*Unknown. 2003. Andes to Amazon botany program at BRIT. Iridos 14: 20-21.
*Valderrama, E., N. Medina, M. Chocce and L. Portal. 2004. Patrones de herbivoría en hojas
jóvenes y adultas de Piper sp. Pages 50-52 in Servat, G. (ed.), unpublished compendium
of student projects from the 2004 field course "Ecología tropical y diseño experimental" at
51
*Vanucci, D. 2003. Erotylidae (Insecta: Coleoptera) en el sotobosque de la cuenca del Río Los
Amigos, Madre de Dios, Perú. Unpublished report for the Asociación para la
Conservación Amazónica. 4 pages.
*Vargas, J. and J. Achicahuala. 2003. Riqueza y abundancia de escarabajos coprofagos
(Coleoptera: Scarabaeidae) en dos tipos de bosque. Pages 122-126 in Servat, G., M.
Rodríguez, D. Cadena, K. Balta, and C. García-Robledo (eds.), unpublished
2003-13," sponsored by the Universidad Nacional de la Amazonía Peruana, Centro
Amazónico de Educación Ambiental e Investigación & the Organization for Tropical
Studies.
*von May, R. 2003. Tercera evaluación de la herpetofauna en el Centro Río Amigos, Madre de
Dios, Perú. Unpublished report for the Asociación para la Conservación Amazónica. 16
pages.
*von May, R. 2004. Geographic distribution, Leptodactylus stenodema. Herpetological Review
35(3): 282.
*von May, R. 2004. Geographic distribution, Osteocephalus buckleyi. Herpetological Review
35(3): 283.
*von May, R. and L. O. Rodríguez. 2004. Guía fotográfica para los anfibios y reptiles del Centro
Río Los Amigos, Madre de Dios, Perú. Unpublished set of color laminas for the
Asociación para la Conservación de la Cuenca Amazónica.
Vriesendorp, C., N. Pitman, R. Foster, I. Mesones & M. Ríos. 2004. Plants/Plantas. Pages 54-61,
141-147 in Pitman, N., R.C. Smith, C. Vriesendorp, D. Moskovits, R. Piana, G. Knell & T.
Wachter (eds.), Perú: Ampiyacu, Apayacu, Yaguas, Medio Putumayo. Rapid Biological
Inventories Report 12. The Field Museum of Natural History, Chicago.
Vriesendorp, C., N. Pitman, J. I. Rojas Moscoso, L. Rivera Chávez, L. Calixto Méndez, M. Vela
Collantes, P. Fasabi Rimachi (eds.). 2005. Perú: Matsés. Rapid Biological Inventories
Report 16. The Field Museum of Natural History, Chicago. 336 pages.
Webb, C. O. 2005. Engineering hope. Conservation Biology 19: 1-3.
*WI. 2004. Ecological applications of multispectral 3D aerial imagery in the Los Amigos Reserve
in eastern Peru. Unpublished grant report by Winrock International (WI) for the Amazon
Conservation Association and the Gordon and Betty Moore Foundation. 14 pages.
*Yánez, I. and M. Gómez. 2004. Rugosidad de los árboles y establecimiento de las epífitas y
hemiepífitas en terraza baja y alta. Pages 134-137 in Servat, G., D. Cadena, K. Balta,
*Yánez, I., M. Gómez, K. Dexter, A. Tauro, R. A. Saldaña, P. Stevenson and M. Espinoza. 2004.
Variación de la densidad y la riqueza de plantas entre claros de diferente edad en un
bosque tropical. Pages 100-104 in Servat, G., D. Cadena, K. Balta, and C. García-
*Zenteno, L., C. González and S. Ríos. 2003. Dime quién te come y te diré quién eres: Herbivoría
en un gradiente de sucesión vegetal. Pages 139-141 in Servat, G., M. Rodríguez, D.
Cadena, K. Balta, and C. García-Robledo (eds.), unpublished compendium of student
projects from OTS course "Ecología de ecosistemas amazónicos 2003-13," sponsored
by the Universidad Nacional de la Amazonía Peruana, Centro Amazónico de Educación
52
Appendix 1.
A LIST OF RECOMMENDED TEXTS
FOR BIOLOGISTS WORKING IN MADRE DE DIOS
compiled by N. Pitman and M. Silman, May 2005; please send comments to

npitman@amazonconservation.org and silmanmr@wfu.edu
LANDSCAPE AND BACKGROUND

• History: Excerpts from Armentia, N. 1887. Navegación del Madre de Dios: Viaje
del Padre Nicolás Armentia. Impresora de La Paz, La Paz.
• Excerpts from Maldonado, J. A. 1899. Relación de la jornada y descubrimiento del
río Manu (hoy Madre de Dios) en 1567. Madrid.
• Hydrology: Excerpts from Goulding, M., C. Cañas, R. Barthem, B. Forsberg and H.
Ortega. 2003. Amazon headwaters: Rivers, wildlife, and conservation in
southeastern Peru. Grafica Biblos S.A., Lima.
• Excerpts from Kalliola, R., and M. Puhakka. 1993. Geografía de la selva baja
peruana. Pages 9-21 in R. Kalliola, M. Puhakka, and W. Danjoy (eds.),
Amazonía peruana: Vegetación húmeda tropical en el llano subandino. PAUT
and ONERN, Jyväskylä, Finland.
• Geology: Excerpts from Palacios M., O., O. Molina G., A. Galloso C., and C.
Reyna L. 1996. Geología de los cuadrángulos de Puerto Luz, Colorado,
Laberinto, Maldonado, Quincemil, Masuco, Astillero y Tambopata. INGEMMET,
Lima.
• Räsänen, M., R. Neller, J. Salo, and H. Jungner. 1992. Recent and ancient fluvial
deposition systems in the Amazonian foreland basin, Peru. Geological Magazine
129:293-306.
• Soils: Osher, L. J. and S. W. Buol. 1998. Relationship of soil properties to parent
material and landscape position in eastern Madre de Dios, Peru. Geoderma
83(1-2): 143-166.
• Mazer, S. J. 1997. Floristic composition, soil quality, litter accumulation, and
decomposition in terra firme and floodplain habitats near Pakitza, Peru. Pages
89-126 in D. E. Wilson and A. Sandoval (eds.), Manu: The biodiversity of
southeastern Peru. Smithsonian Institution and Editorial Horizonte, Lima.
• A colpa paper – which?
• Climate: For regional context one of the basin-wide climate papers, maybe Jose
Marengo’s book chapter?
• A paleoclimate paper by Baker et al. or Bush et al.
• Vanschaik, C. P., J. W. Terborgh and S. J. Wright. 1993. The phenology of tropical
forests: Adaptive significance and consequences for primary consumers. Annual
Review of Ecology and Systematics 24: 353-377.
• Foster, M. S. and J. Terborgh. 1998. Impact of a rare storm event on an
Amazonian forest. Biotropica 30(3): 470-474.
AQUATIC ECOSYSTEMS
• Excerpts from Goulding, M., C. Cañas, R. Barthem, B. Forsberg and H. Ortega.
2003. Amazon headwaters: Rivers, wildlife, and conservation in southeastern
Peru. Grafica Biblos S.A., Lima.
• Excerpts from Barthem, R., M. Goulding, B. Forsberg, C. Cañas and H. Ortega.
2003. Aquatic ecology of the Río Madre de Dios: Scientific bases for Andes-
Amazon headwaters conservation. Asociación para la Conservación de la
53
Cuenca Amazónica/Amazon Conservation Association, Lima.

• Ortega, H. 2003. Peces del sector peruano del corredor de conservación
Vilcabamba-Amboró. Unpublished report of the Critical Ecosystem Partnership
Fund and Conservation International.
• Schenck, C., J. Groenendijk, F. Hajek, E. Staib and K. Frank. 2003. Giant otters in
the Peruvian rainforest: Linking protected area conditions to species needs.
Pages 341-357 in J. A. Bissonette and I. Storch (eds.), Landscape ecology and
resource management: Linking theory with practice. Island Press, Washington,
DC.
• Excerpts from Schenck, C. 1999a. Lobo de río (Pteronura brasiliensis): Presencia,
uso del hábitat y protección en el Perú. GTZ & INRENA. Lima. 176 pages.
PLANTS
• Regional context: Gentry, A. H. 1997. Lowlands of Manu National Park: Cocha
Cashu Biological Station, Peru. Pages 360-363 in S. D. Davis, V. H. Heywood,
O. Herrera-MacBryde, J. Villa-Lobos and A. C. Hamilton (eds.), Centres of plant
diversity: A guide and strategy for their conservation. Volume 3. WWF and IUCN,
Cambridge, UK.
• Terborgh, J. and E. Andresen. 1998. The composition of Amazonian forests:
patterns at local and regional scales. Journal of Tropical Ecology 14: 645-664.
• Excerpts from ter Steege, H., N. Pitman, D. Sabatier, H. Castellanos, P. Van der
Hout, D. C. Daly, M. Silveira, O. Phillips, R. Vasquez, T. Van Andel, J.
Duivenvoorden, A. A. De Oliveira, R. Ek, R. Lilwah, R. Thomas, J. Van Essen, C.
Baider, P. Maas, S. Mori, J. Terborgh, P. N. Vargas, H. Mogollon and W.
Morawetz. 2003. A spatial model of tree alpha-diversity and tree density for the
Amazon. Biodiversity and Conservation 12(11): 2255-2277.
• Gentry, A. H. and R. Ortiz. 1993. Patrones de composición florística en la
Amazonía peruana. Pages 155-166 in R. Kalliola, M. Puhakka and W. Danjoy
(eds.), Amazonía peruana: Vegetación húmeda tropical en el llano subandino.
Proyecto Amazonia of the Universidad de Turku (PAUT), and Oficina Nacional
de Evaluación de Recursos Naturales (ONERN), Jyväskylä.
• Swamps: Excerpts from Ancaya, E. J. 2002. Floristics and hydrology of upper
Amazonian swamps. Master's thesis. Department of Biology, Wake Forest
University, Winston-Salem, USA.
• Floodplains: Foster, R. B. 1990. The floristic composition of the Rio Manu
floodplain forest. Pages 99-111 in A. H. Gentry (ed.), Four Neotropical
rainforests. Yale University Press, New Haven, USA.
• Gentry, A. H. and J. Terborgh. 1990. Composition and dynamics of the Cocha
Cashu mature floodplain forest, Peru. Pages 542-564 in A. H. Gentry (ed.), Four
Neotropical rainforests. Yale University Press, New Haven.
• Terborgh, J. and K. Petren. 1991. Development of habitat structure through
succession in an Amazonian floodplain forest. Pages 28-46 in S. Bell, E. D.
McCoy and H. R. Mushinsky (eds.), Habitat structure: The physical arrangement
of objects in space. Chapman and Hall, New York.
• Uplands: Pitman, N. C. A., J. W. Terborgh, M. R. Silman, P. Nunez, D. A. Neill, C.
E. Ceron, W. A. Palacios and M. Aulestia. 2001. Dominance and distribution of
tree species in upper Amazonian terra firme forests. Ecology 82(8): 2101-2117.
• Bamboo: Griscom, B. W. and P. M. S. Ashton. 2003. Bamboo control of forest
succession: Guadua sarcocarpa in Southeastern Peru. Forest Ecology and
Management 175(1-3): 445-454.
54
• Silman, M. R., E. J. Ancaya and J. Brinson. 2003. Los bosques de bambú en la

Amazonía occidental. Pages 63-74 in R. Leite Pitman, N. Pitman and P. Álvarez
(eds.), Alto Purús: Biodiversidad, conservación y manejo. Center for Tropical
Conservation, Lima.
• Brazil nuts: Excerpts from Cornejo Valverde, F. 2003. Historia natural de la
castaña (Bertholletia excelsa Humb. & Bonpl.) y propuestas para su manejo.
Asociación para la Conservación de la Cuenca Amazónica, Puerto Maldonado.
52 pages.
• Excerpts from Ortiz, E. 2002. Brazil nut (Bertholletia excelsa). Pages 61-74 in P.
Shanley, A. Pierce, S. Laird, and A. Guillen (eds.), Tapping the green market:
Certification and management of non-timber forests products. People and Plants
Conservation Series, UNESCO.
INSECTS
• Excerpts from Erwin, T. L. 1991. Natural history of the carabid beetles at the
BIOLAT Biological Station, Rio Manu, Pakitza, Peru. Revista Peruana de
Entomología 33: 1-85.
• Lamas, G. 1997. Comparing the butterfly faunas of Pakitza and Tambopata, Madre
de Dios, Peru, or Why is Peru such a megadiversity country. Pages 165-168 in
H. Ulrich (ed.), Tropical diversity and systematics: Proceedings of the
International Symposium on Biodiversity and Systematics in Tropical
Ecosystems. Zoologische Forchunginstitut und Museum Alexander Koenig,
Bonn.
• Pearson, D. L. and J. A. Derr. 1986. Seasonal patterns of lowland forest floor
arthropod abundance in southeastern Peru. Biotropica 18(3): 244-256.
• Tobin, J. E. 1997. Competition and coexistence of ants in a small patch of
rainforest canopy in Peruvian Amazonia. Journal of the New York Entomological
Society 105(1-2): 105-112.
• Wilson, E. O. 1987. The arboreal ant fauna of Peruvian Amazon forests: A first
assessment. Biotropica 19(3): 245-251.
• Yu, D. W. and N. E. Pierce. 1998. A castration parasite of an ant-plant mutualism.
Proceedings of the Royal Society of London Series B-Biological Sciences
265(1394): 375-382.
AMPHIBIANS AND REPTILES

• Cadle, J., J. Icochea, J. Pérez Zúñiga, A. Portilla and C. Rivera. 2002. La
herpetofauna encontrada en el Refugio Juliaca y en el Puesto de Vigilancia
Enahuipa del Santuario Nacional Pampas del Heath. Pages 52-57 in J. R.
Montambault (ed.), Informes de las evaluaciones biológicas Pampas del Heath,
Perú; Alto Madidi, Bolivia; y Pando, Bolivia. Conservation International Bulletin of
Biological Assessment 24, Washington, DC, USA.
• Rodríguez, L. O. 1990. A preliminary overview of the herpetofauna at Cocha
Cashu, Manu. Pages 410-425 in A. H. Gentry (ed.), Four Neotropical rainforests.
Yale University Press, New Haven, or Rodríguez, L. O. 1992. Structure et
organisation du peuplement d’anoures de Cocha Cashu, Parque National Manu,
Amazonie Péruvienne. Terre et Vie 47(2): 151-197.
• von May, R. 2003. Tercera evaluación de la herpetofauna en el Centro Río
Amigos, Madre de Dios, Perú. Unpublished report for the Asociación para la
Conservación Amazónica. 16 pages.
55
BIRDS
• Kratter, A. W. 1997. Bamboo specialization by Amazonian birds. Biotropica 29(1):
100-110.
• Parker, T. A., III, P. K. Donahue and T. S. Schulenberg. 1994. Birds of the
Tambopata Reserve (Explorers' Inn). Pages in R. B. Foster, J. L. Carr and A. B.
Forsyth (eds.), The Tambopata-Candamo Reserved Zone of southeastern Peru:
A biological assessment. Conservation International, Washington, D. C.
• Terborgh, J., S. K. Robinson, T. A. Parker, III, C. A. Munn and N. Pierpont. 1990.
Structure and organization of an Amazonian forest bird community. Ecological
Monographs 60(2): 213-238.
• Excerpts from the old volume of Monographs in Ornithology that has some classic
EBCC papers on mixed flocks, etc. Scott R’s work and Fitzpatrick’s stuff on
flycatchers.
MAMMALS
• Regional context: Voss, R. S. and L. H. Emmons. 1996. Mammalian diversity in
Neotropical lowland rainforests: a preliminary assessment. Bulletin of the
American Museum of Natural History 230(26): 1-115.
• Emmons, L. H. 1987. Comparative feeding ecology of felids in a Neotropical
rainforest. Behavioral Ecology and Sociobiology 20(4): 271-283.
• Kiltie, R. A. and J. Terborgh. 1983. Observations on the behavior of rain forest
peccaries in Peru: Why do white-lipped peccaries form herds? Zeitschrift fur
Tierpsychologie 62(3): 241-255.
• Leite Pitman, R., H. Beck and P. M. Velazco. 2003. Mamíferos terrestres y
arbóreos de la selva baja de la Amazonía peruana: Entre los ríos Manu y Alto
Purús. Pages 109-124 in R. Leite Pitman, N. Pitman and P. Álvarez (eds.), Alto
Purús: Biodiversidad, conservación y manejo. Center for Tropical Conservation,
Lima.
• Excerpts from Terborgh, J. 1983. Five New World primates: A study in
comparative ecology. Princeton University Press, Princeton, USA.
• Abstracts/excerpts from other classic primate studies at Cocha Cashu, e.g.,
Wright, P. C. 1994. The behavior and ecology of the owl monkey. Pages 97-112
in J. Baer, R. Weller and I. Kakomo (eds.), Aotus: The owl monkey. Academic
Press, San Diego, Anne Goldizen on tamarins.
PLANT-ANIMAL INTERACTIONS
• Silman, M. R., J. W. Terborgh, and R. A. Kiltie. 2003. Population regulation of a
dominant-rain forest tree by a major seed-predator. Ecology 84:431-438.
• Terborgh, J. 1986. Keystone plant resources in the tropical forest. Pages 330-344
in M. E. Soulé (ed.), Conservation biology: The science of scarcity and diversity.
Sinauer Associates, Sunderland, USA.
• Wright, S. J., M. E. Gompper and B. Deleon. 1994. Are large predators keystone
species in neotropical forests? The evidence from Barro Colorado Island. Oikos
71(2): 279-294.
CONSERVATION
• Álvarez, N. L. and L. Naughton-Treves. 2003. Linking national agrarian policy to
deforestation in the Peruvian Amazon: A case study of Tambopata, 1986-1997.
Ambio 32(4): 269-274.
• Dourojeanni, M. J. 2003. Impactos socioambientales de las carreteras
56
transfronterizas y fronterizas en los departamentos de Madre de Dios y Ucayali y

capacidad de respuesta del Perú. Pages 237-246 in R. Leite Pitman, N. Pitman
and P. Álvarez (eds.), Alto Purús: Biodiversidad, conservación y manejo. Center
for Tropical Conservation, Lima.
• Escobal, J. and U. Aldana. 2003. Are nontimber forest products the antidote to
rainforest degradation? Brazil nut extraction in Madre De Dios, Peru. World
Development 31(11): 1873-1887.
• Peres, C. A., C. Baider, P. A. Zuidema, L. H. O. Wadt, K. A. Kainer, D. A. P.
Gomes-Silva, R. P. Salomao, L. L. Simoes, E. R. N. Franciosi, F. C. Valverde, R.
Gribel, G. H. Shepard, M. Kanashiro, P. Coventry, D. W. Yu, A. R. Watkinson
and R. P. Freckleton. 2003. Demographic threats to the sustainability of Brazil
nut exploitation. Science 302(5653): 2112-2114.
• Rodríguez, L. O., and K. R. Young. 2000. Biological diversity of Peru: Determining
priority areas for conservation. Ambio 29(6): 329-337.
• Yu, D. W., T. Hendrickson and A. Castillo. 1997. Ecotourism and conservation in
Amazonian Peru: Short-term and long-term challenges. Environmental
Conservation 24(2): 130-138.
• Excerpts from the Los Amigos Conservation Concession and Manu National Park
management plans.
PEOPLE
• Clark, C., L. Michael, and C. Beier. 2005. The Mashco Piro of the Río Purús
region: Factors affecting their well-being and self-determination in a changing
world. Unpublished report of the Cabeceras Aid Project. 29 pages.
• Excerpts from Huertas, Beatriz. 2002. Los pueblos indígenas en aislamiento: Su
lucha por la sobrevivencia y la libertad. IWGIA, Lima.
• IIAP. 2002. Propuesta de zonificación ecológica económica de la región Madre de
Dios: Versión corregida en base a los acuerdos de la asamblea regional sobre
Zonificación Ecológica Económica (ZEE). Unpublished report of the Instituto de
Investigaciones de la Amazonía Peruana (IIAP).
• Phillips, O., A. H. Gentry, C. Reynel, P. Wilkin and B. C. Galvez Durand. 1994.
Quantitative ethnobotany and Amazonian conservation. Conservation Biology
8(1): 225-248.
• Shepard, G. H., Jr. 2001. Rainforest habitat classification among the Matsigenka of
the Peruvian Amazon. Journal of Ethnobiology 21(1): 1-38.
OTHER
• D'Achille, B. Excerpts from Uturunkusuyo: El territorio del jaguar. Lima.
• Leopold, A. [1924] 1991. The River of the Mother of God. Essay from The river of
the mother of God and other essays. University of Wisconsin Press, Madison,
USA.
• Symington, M. xx de julio. Pages xx-xx in M. B. Mulder and W. Logsdon (eds.),
I’ve been gone far too long: Field study fiascoes and expedition disasters. RDR
Books.
• Terborgh, J. 2000. In the company of humans. Natural History 109: 54-63.
57
Appendix 2.
RESEARCH PROJECTS ACTIVE AT LOS AMIGOS, 2000-2006
ordered by topic, then alphabetically

by surname of lead investigator
CLIMATE, GEOLOGY, SOILS, AND HYDROLOGY

Geology of the Los Amigos area
Dr. Pierre-Olivier Antoine (France), Dr. Patrice Baby (France), Dr. Stephane Brusset
(France), Dr. Martin Roddaz (France), Wilber Hermoza (Peru)
Institut de Recherche pour le Developpement (France)
2003
Geology of the Cerro Colorado cliff

Dr. Kenneth Campbell, Jr. (USA), Dr. Lidia Romero-Pittman (Peru)
Natural History Museum of Los Angeles County (USA), Instituto Geológico, Minero y
Metalúrgico (Peru)
2006
Global patterns of soil biodiversity: Implications for ecosystem function

Dr. Edward Ayres (USA), Dr. Richard Bardgett (UK), Dr. Johnson Nkem (Cameroon), Dr.
Diana Wall (USA)
Colorado State University (USA), Lancaster University (UK)
2005
Hydrology of the Madre de Dios watershed

Dr. Michael Goulding (USA), Carlos Cañas (Peru), Rolando Barthem (Brazil), and others
Amazon Conservation Association and other institutions
2001-2003
Geomorphology and soil development in the Los Amigos area

Lars Jespersen (Denmark), Kristian Løkke Kristensen (Denmark), Christian Bjergager
(Denmark), and Dr. Lars Rasmussen (Denmark)
Royal Veterinary and Agricultural University
2005
A GIS study of landslides in the lower Los Amigos foreland of Peru

Suresh Khanal (Nepal)
Texas Christian University (USA)
2005-2006
A reconnaissance of the transient storage properties and nitrogen dynamics of streams

in the Los Amigos Concession
Dr. Jim McNamara (USA)
Boise State University (USA)
2004
Monitoring climatic conditions of palm swamps with a micro-sensor network

William Mantzell (USA)
58
Rice University
2004
The River Continuity Concept and its applicability to Neotropical river systems
Eduardo Oyague (Peru)
Universidad Nacional Agraria La Molina (Peru)
2004-2005
Chemical composition of colpa soils

Dr. Lars Rasmussen (Denmark)
Royal Veterinary and Agricultural University
2004
Exploratory research on Quaternary sedimentary history of fluvial terraces in the Madre

de Dios basin
Dr. Catherine Rigsby (USA), Dr. Paul Baker (USA), Erin Hemric (USA)
East Carolina University (USA), Duke University (USA)
2005
Reconaissance of Madre de Dios streams

Dr. Bernard Sweeney (USA), Ing. Giancarlo Barbieri (Peru), Dr. Ralph Flowers (USA),
Dr. David Funk (USA), Dr. Michael Gentile (USA), Dr. Jan Surma (USA)
Stroud Water Research Center (USA)
2006
VEGETATION
Inventory of Sapindaceae in the Los Amigos concession
Dr. Pedro Acevedo (USA)
Smithsonian Institution (USA)
2004
Inventory of commercially important tree species in the logging concessions north of

CICRA
Ing. Jorge Alva (Peru), Ing. Percy Núñez (Peru), Dr. Nigel Pitman (USA)
Asociación para la Conservación de la Cuenca Amazónica (Peru)
2004-2005
Quantitative inventory of upland and floodplain tree plots

Dante Anton (Peru), Ing. Fernando Cornejo (Peru), Therany Gonzáles (Peru), Dr. John
Janovec (USA), Piher Maceda (Peru), Ing. Percy Núñez (Peru), Dr. Nigel Pitman (USA),
Dr. John Terborgh (USA), Mathias Tobler (Switzerland), César Vela (Peru), and others
Amazon Conservation Association (USA), Botanical Research Institute of Texas (USA),
Duke University (USA), Universidad Nacional Agraria La Molina (Peru), Universidad
Nacional San Antonio Abad del Cusco (Peru), and others
(1993-) 2002-2006
Spatial distribution of trees with different dispersal modes

Cecilia Blundo (Argentina) and Alejandra Tauro (Argentina)
Universidad de Córdoba (Argentina)
2005
59
Lianas at Los Amigos: Dominants, distribution, and habitat preferences

Dr. Robyn Burnham (USA)
University of Michigan (USA)
2006
Distribution of lichens in the Los Amigos Conservation Concession

Jorge Carrillo (Peru)
Universidad Peruana Cayetano Heredia (Peru)
2004
Taxonomy and diversity of pteridophytes in different vegetation types of the Los Amigos
watershed
Benjamín Chambi (Peru)
Universidad Nacional San Antonio Abad del Cusco - Puerto Maldonado (Peru)
2005-2006
Genetic relationships and densities of Hedyosmum (Chloranthaceae) in montane and

lowland forests of southeastern Peru
Carmen Chávez (Peru)
Wake Forest University (USA)
2004
Inventory of orchids
Miguel Chocce (Peru)
Universidad Nacional Mayor San Marcos (Peru)
2003
Inventory and harvest potential of sangre de grado and copaiba at CICRA

Mickelly Cuba (Peru)
2003-2004
Seed-fungal interactions of pioneer tree species in tropical forests

Dr. Jim Dalling (UK), Marcos Ríos (Peru)
University of Illinois (USA)
2006
Conservation prioritization of orchids in the Los Amigos Concession

Miluzka Damian (Peru)
Universidad Ricardo Palma (Peru)
2005
Genetic diversity in Cedrela odorata populations in Madre de Dios

Amanda De La Torre (Peru)
2003
Ecology and geographical distribution of Inga in Amazonian Peru

Kyle Dexter (USA)
Duke University (USA)
2004-2006
60
12-year recensus of a 2-ha permanent plot in terra firme forest near CICRA
Rolando Díaz (Peru) and others
Universidad Nacional San Antonio Abad del Cusco - Puerto Maldonado (Peru)
2005
Long-term monitoring of leaf area index in two forest types at CICRA

Dr. Marc Dubois (France) and Dr. Louise Emmons (USA)
CEA Saclay, Service de Physique de l’Etat Condensé (France) and Smithsonian
Institution (USA)
2002-2006
Influence of light, soil moisture, and soil fertility in the early growth of Cedrela spp.
Ing. César Flores (Peru)
Yale School of Forestry (USA)
2001-2002
Inventory of Brazil nut trees in a 12-km2 area on the CICRA terrace

Ing. César Flores (Peru) and others
Yale School of Forestry (USA)
2002
Preliminary inventory of the vegetation of Los Amigos

Dr. Robin Foster (USA), Ing. Hamilton Beltrán (Peru), and others
Field Museum (USA) and Museo de Historia Natural de UNMSM (Peru)
2001
Linking tree alpha-diversity to environmental gradients and remote sensing in western

Peruvian Amazonia
Roosevelt García (Peru), Nallarett Dávila (Peru), Karla Meza (Peru), Marcos Ríos (Peru)
University of Missouri-St. Louis (USA)
2004
Influence of tree-fall gaps on the natural regeneration of five economically important

timber species in the Los Amigos watershed
Therany Gonzales (Peru)
Universidad Nacional San Antonio Abad del Cusco-Puerto Maldonado (Peru)
2004
Vegetation mapping along 170 km of trails in and around the Los Amigos Conservation
Concession
Therany Gonzales (Peru)
Universidad Nacional San Antonio Abad del Cusco-Puerto Maldonado
2005-2006
Structure and dynamics of Mauritia flexuosa populations in the Los Amigos Conservation
Concession
Nelson Gutiérrez (Peru)
Université de Toulouse (France)
2004-2005
61
Vegetation types and hydrology of the Los Amigos area

Dr. Steve Hamilton (USA)
Michigan State University (USA)
2004
Ecophysiology of carbon uptake and water use by the palm Mauritia flexuosa L.
Viviana Horna (Peru), Reiner Zimmermann (Germany)
University of Goettingen (Germany)
2005-2006
Natural history, population biology, and pollination biology of vanilla orchids (Vanilla
spp.)
Ethan Householder (USA), Alex Van Dam (USA)
Texas Christian University (USA), University of California-Riverside (USA)
2005-2006
Los Amigos botany project

Dr. John Janovec (USA), Ing. Fernando Cornejo (Peru), Piher Maceda (Peru), Pedro
Maceda (Peru), Angel Balarezo (Peru) and others
Botanical Research Institute of Texas (USA)
2001-2006
The distribution, abundance, clump characteristics and techniques for managing Guadua
cf. angustifolia, Bambuseae, a potential non-wood forest product, in Madre de Dios,
Peru
Chris Kirkby (UK)
University of York (UK)
2001-2002
Inventory of mosses and lichens

Dr. Piers Majestyk (USA)
New York Botanical Garden (USA)
2002
Surveying vegetation with high-resolution aerial photographs in the Los Amigos

Conservation Concession
Josué Mercado (Peru)
Universidad Nacional San Antonio Abad del Cusco (Peru)
2006
World herbivory project

Dr. Angela Moles (New Zealand), Therany Gonzales (Peru)
Macquarie University (Australia)
2005-2006
Los Amigos jungle cucumbers: A study of five sympatric Gurania species

Dr. Amanda Neill (USA)
Botanical Research Institute of Texas (USA)
2001-2002
Ecology and dynamics of bamboo
62
Jean Olivier (France), Sam Dubois (France)

Université de Toulouse (France)
2002-2005
Seedling recruitment and resource availability in the western Amazon

Timothy Paine (USA)
Louisiana State University (USA)
2005-2006
Light levels and Brazil nut regeneration

Ing. Ronald Rado Quispe (Peru)
Universidad Nacional San Antonio Abad del Cusco-Puerto Maldonado (Peru)
2001-2004
Effects of diminished seed disperser abundances on tree recruitment in tierra firme

forests: A comparison of the spacing dynamics of the canopy tree communities of
pristine vs. impacted sites
Varun Swamy (India)
Duke University (USA)
2005-2006
Structure, floristic composition, and ecology of epiphytic plant communities

Michael Vega (Peru)
2004
PLANT-ANIMAL INTERACTIONS
Impacts of peccaries on plant communities in Los Amigos and Cocha Cashu
Dr. Harald Beck (Germany), Karim Ledesma (Peru)
Towson University (USA)
2004-2006
Wood-boring cerambycid beetles associated with the tree family Lecythidaceae

Dr. Amy Berkov (USA), Pedro Centeno (Peru), Eulogio Condori (Peru)
City College of New York (USA)
2003-2006
Fauna associated with the bamboo Guadua weberbaueri Pilger

Patricia Feria (Mexico), Jon Dale (USA)
2006
Seed dispersal by the yellow-footed tortoise, Geochelone denticulata, in Madre de Dios,

Peru
Adriana Guzman (Colombia)
Universidad de Los Andes (Colombia)
2006
63
Ecology of birds specializing on bamboo forest

Dan Lebbin (USA)
Cornell University (USA)
2002-2004
Area requirements of Amazonian keystone species

Dr. George Powell (USA), Krista Adamek (Canada), Thomas Adamek (Canada), Dra.
Renata Leite Pitman (Brazil), Rafael Mares (Panama), Sue Palminteri (USA), Raul
Tupayachi (Peru), and others
World Wildlife Fund USA and other institutions
2003-2006
Habitat fragmentation and ecology of bird specialists on bamboo

Wendy Tori (Peru)
2003
INVERTEBRATES
The evolution of social parasitism in Megalomyrmex ants
Rachelle Adams (USA)
University of Texas (USA)
2004
Long-term monitoring of butterfly and dung beetle communities at Los Amigos

Dr. Phil DeVries (USA), Trond Larsen (USA), Pedro Centeno (Peru), Eulogio Condori
(Peru)
University of New Orleans (USA), Princeton University (USA)
2003-2005
Variation in the spider community during dry and wet seasons in the Los Amigos
watershed
Mariajosé Deza (Peru)
2005-2006
Diversity, composition, distribution, and food plant use of ants in understory vegetation in
Amazonian Peru
Megan Frederickson (USA)
Stanford University (USA)
2005
Assessing diversity and spatial organization of Neotropical drosophilids: The canopy as

unexplored territory
Craig Gibbs (USA)
City University of New York (USA)
2004-2005
Biodiversity of the Arctiinae (Lepidoptera: Arctiidae) in the Los Amigos Conservation

Concession
Juan Grados (Peru), Angélico Asenjo (Peru)
64
Museo de Historia Natural (Peru)

2006
Inventory of Mollusca assemblages in the Los Amigos watershed

Cristian Ituarte (Argentina), Gabriela Cuezzo (Argentina), Rina Ramírez (Perú)
Museo de la Plata (Argentina), Universidad Nacional de Tucumán (Argentina),
Universidad Nacional Mayor de San Marcos (Perú)
2006
Ground beetles in bamboo forests: Community patterns and patch specificity in a

heterogeneous landscape
Jenny Jacobs (USA)
San Francisco State University (USA)
2006
Biology of Apterostigma ants

Stephanie Johnson (USA)
University of Maryland (USA)
2005
Ecology of dung beetles

Trond Larsen (USA)
Princeton University (USA)
2000-2005
Evolutionary ecology of the Attine ant-microbe mutualism

Ainslie Little (USA), Dr. Matías Cafaro (Argentina)
University of Wisconsin (USA)
2004
Inventory of Cyclopoida copepods

Mabel López (Peru)
Universidad Nacional Mayor de San Marcos (Peru)
2006
Survey of leaf-litter ants and dung beetles at CICRA

Dr. Chris Marshall (USA), Dr. Ted Schultz (USA), Jeffrey Sosa-Calvo (Colombia),
Smithsonian Institution (USA), University of Maryland (USA)
2004
Systematics and distribution of scorpions in the Madre de Dios watershed

Dr. José Antonio Ochoa (Peru), Juan Carlos Chaparro (Peru)
2005
Inventory of Diptera
Steven Paiero (Canada), John Klymko (Canada)
University of Guelph (Canada)
2006
Evolution of host plant use in the butterfly subfamily Satyrinae (Lepidoptera:

Nymphalidae)
65
Carlos Peña (Peru)

Stockholm University (Sweden)
2005
A taxonomic revision of the Neotropical non-leafcutting fungus-growing ant genus

Mycocepurus
Christian Rabeling (Germany)
Eberhard-Karls University Tübingen (Germany)
2004
Inventory of meliponnine and euglossine bees

Dr. David Roubik (USA)
Smithsonian Tropical Research Institute (USA)
2003
Molecular phylogeny of ants in the genus Pheidole

Corrie Saux (USA)
Harvard University (USA)
2004
Testing hypotheses of tropical diversification in leafcutter ants

Scott Solomon (USA)
University of Texas (USA)
2004
Revision of the Neotropical primitive fungus-growing ants in an Amazonian forest

Jeffrey Sosa-Calvo (Colombia)
University of Maryland (USA)
2004-2005
Inventory of erotylid beetles and their fungal hosts

Dino Vanucci (Peru)
2003
Behavioral and chemical ecology of Neotropical ant gardens

Elsa Youngsteadt (USA)
North Carolina State University (USA)
2004-2005
FISH
Ichthyological inventory of the Madre de Dios and Los Amigos rivers
Dr. Michael Goulding (USA), Carlos Cañas (Peru), Dr. Rolando Barthem (Brazil), and
others
Amazon Conservation Association (USA), Asociación para la Conservación de la
Cuenca Amazónica (Peru), and other institutions
2001-2003
Ichthyological inventory of the Amiguillos River

Max Hidalgo (Peru), Dr. Hernán Ortega (Peru), Soraya Martínez (Peru), Vanessa
66
Chipollini (Peru), and others

Museo de Historia Natural UNMSM (Peru), Universidad Ricardo Palma (Peru),
2004-2006
AMPHIBIANS AND REPTILES

Long-term monitoring of fauna along the Los Amigos River
Hernán Collado (Peru), Wilberth Concha (Peru), Jerry Martínez (Peru), Alberto
Melendez (Peru), Jorge Pérez (Peru), Fernando Pinto (Peru), Edwin Quispe (Peru),
David Ttito (Peru), Eriberto Torres (Peru), Dr. Nigel Pitman (USA)
2004-2006
Elevated exposure to airborne pollutants in arboreal frog larvae of the neotropics

Dante Fenolio (USA), Jim Stout (USA)
University of Miami (USA)
2006
Reproductive ecology of Hyla frogs in the leucophyllata group

Giuseppe Gagliardi (Peru)
Universidad Nacional de la Amazonía Peruana (Peru)
2004-2005
Seed dispersal by the yellow-footed tortoise, Geochelone denticulata, in Madre de Dios,

Peru
Adriana Guzman (Colombia)
Universidad de Los Andes (Colombia)
2006
Status and management of the river turtle, Podocnemis unifilis, in the Los Amigos River
Ana María Mansilla (Peru), David Ttito (Peru), Edwin Quispe (Peru)
2004-2006
Cannibalism in tadpoles of Dendrobates biolat

Eva Medina (Peru)
2003
Ecology of microteiid lizards (Gimnophthalmidae) at CICRA

Juana Suárez (Peru)
University Nacional Mayor de San Marcos (Peru)
2004-2005
Inventory of the Los Amigos herpetofauna

Rudolf von May (Peru)
2001-2003
Anuran beta diversity across forest types in a lowland Amazonian rainforest
67
Rudolf von May (Peru)

Florida International University (USA)
2005-2006
BIRDS
2004-2006
Ecology of birds specializing on bamboo forest

Dan Lebbin (USA)
2002-2005

2003-2006
Preliminary study of the behavior and ecology of the warbling antbirds Hypocnemis
cantator peruviana and Hypocnemis cantator collins
Dr. Nathalie Tobias (UK), Dr. Joe Tobias (UK), Paulo Pulgarín (Colombia)
Cambridge University (UK)
2004-2006
Inventory of the Los Amigos avifauna

Dr. Douglas Stotz (USA), Tatiana Pequeno (Peru)
Field Museum (USA), Museo de Historia Natural UNMSM (Peru)
2002
Habitat fragmentation and ecology of bird specialists on bamboo

Wendy Tori (Peru)
2003
Ecological roles and habitat use of two species of forest-falcons in the Amazonian
rainforest
Ursula Valdez (Peru), Peggy Shrum (USA), Raul Báez (Peru), Laura Riba Hernández
(Costa Rica), and others
University of Washington (USA)
2004-2006
Ecology of Tachycineta swallows across the Americas

Dr. David Winkler (USA) and others
68
2003-2004
MAMMALS
Ecology and behavior of the emperor tamarin, Saguinus imperator
Israel Aragón (Peru)
Universidad Nacional San Antonio Abad del Cusco (Peru)
2003-2004
Frugivorous bats (Phyllostomidae) that visit clay licks in a Neotropical rainforest:

Implications for seed dispersal and plant community structure
Adriana Bravo (Peru)
Louisiana State University (USA)
2005

2004-2006
Phylogeography of Proechimys
Maria de Angelo (USA)
Yale University (USA)
2004
Ecology of mineral licks: Function, effects on vertebrate biomass, and management

implications
Dr. Louise Emmons (USA), Mogens Trolle (Denmark), Dyana La Rosa (Peru)
Smithsonian Institution (USA)
2000-2003
Inventory of giant otters and their habitat in the Los Amigos watershed
Freddy Ferreyra (Peru) and others
Frankfurt Zoological Society (Germany)
2004
Understanding the pair bond in Callicebus brunneus: Male and female interests
Jenna Lawrence (USA) and others
Columbia University (USA)
2002, 2004-2005
Diversity and conservation of bat communities at CICRA

Cristina Lopez Wong (Peru), Jhony Ríos (Peru), Giuseppe Gagliardi (Peru)
Universidad Nacional de la Amazonía Peruana (Peru)
2005-2006
69
2003-2006
Ecology and systematics of marsupials in Neotropical rainforests

Oswaldo Ramírez (Peru), Margarita Arana (Peru), Enrique Bazán (Peru)
Universidad Cayetano Heredia (Peru)
2006
Historical distribution of giant otters in Madre de Dios

Marne Suárez (Peru)
World Wildlife Fund Peru (Peru)
2004
Ecology of tapirs (Tapirus terrestris) in the Los Amigos watershed

Mathias Tobler (Switzerland), Dr. Louise Emmons (USA), Carlos Sánchez (México), Darío Cruz,
Jr. (Peru), Javier Huinga (Peru), and others
Texas A&M University (USA), Botanical Research Institute of Texas (USA), Smithsonian
Insitution (USA)
2002-2006
Vertical stratification of bats in the genus Artibeus (Phyllostomidae) at CICRA

Sandra Velazco (Peru)
Universidad Nacional Mayor San Marcos (Peru)
2005-2006
OTHER
The diversity of the eukaryote microbial groups Alveolata and Cercozoa in a tropical
rainforest environments using microscopy and molecular inference
David Bass (UK), Thomas Richards (UK)
University of Oxford (UK), Natural History Museum London (UK)
2005
Preliminary inventory of the mycoflora at CICRA

Romina Gazis (Peru)
Texas Christian University (USA), Universidad Ricardo Palma (Peru)
2003-2005
Autonomous radio-tracking project

Dr. Robert MacCurdy (USA)
2004
Bibliography of biology or conservation works from Madre de Dios

Dr. Nigel Pitman (USA)
Amazon Conservation Association (USA)
2004-2005
Inventory of polypore fungi at CICRA
70
Alejandro Sánchez Taranov (Peru)

Universidad Cayetano Heredia (Peru)
2006
Ecological applications of multispectral 3D aerial imagery in the Los Amigos reserve in

eastern Peru
Dr. Dana Slaymaker (USA), Dr. Ken MacDicken (USA), Dr. Alfredo Unda (Canada),
Aaron Dushku (USA)
Winrock International (USA)
2003-2005
71
Appendix 3.
COMPANION NOTES FOR THE PRINCIPAL TRAILS
OF THE LOS AMIGOS RESEARCH STATION
The stairs
The station’s infamously unwelcoming approach trail ascends 251 steps to the station
terrace, some 40 m above the river. The oldest person to have made the climb was 75
years old at the time; upon reaching the top she triumphantly lit a cigarette. The
youngest person to have made the climb with no assistance was three years old. She
took her time going up, and so should you.
The forest to either side of the stairs is less than 20 years old; when the station was a
mining camp this hillside was bare, and it was possible to see the port from the head of
the stairs and vice versa. Near the base of the stairs (but not visible from them) is an
overgrown plantation, where a small hillside planted in pineapple overlooks a stand of
cacao, banana, and papaya trees. We are currently deciding whether to reactivate the
plantation or remove it. The argument for reactivating the plantation is that it would
relieve us of the absurd practice of bringing bananas from Puerto Maldonado. The
argument for letting the area go back to nature is that only two forests in the Río Madre
de Dios floodplain are currently protected from development, and this is one of them.
The obvious compromise solution is to restore the plantation to wilderness and buy
bananas from neighboring farms.
The hillside vegetation along the stairs already has a healthy understory of aroids, ferns
and treeferns, Costus, Piper, and melastomes. A native bamboo (apparently Guadua
weberbaueri) is scattered around the first and second descansos and dominates a large
patch just behind the first. A couple of large mango trees at the first descanso and a
smaller citrus tree on the slope below are reminders of the area’s recent past. Several of
the largest trees here belong to the genus Erythrina, a legume characteristic of disturbed
river banks (there is one at the dock in Laberinto) with striking hummingbird-pollinated
red flowers. There are also two large fig trees (Ficus insipida), one at the second
descanso and one at the top of the stairs, as well as some fast-growing youngsters that
will dominate the canopy of this forest in a few years.
Small monkeys are seen relatively frequently from the stairs, particularly emperor
tamarins (Saguinus imperator), saddleback tamarins (Saguinus fuscicollis), and squirrel
monkeys (Saimiri sciureus).
Trocha Aerodromo (1)

This trail is part of an old road built by the gold-mining company that occupied the camp
in the 1980s and 1990s. (The rest of the road is now called Trocha Carretera; see
below.) The road was paved with river stones so that the company could move its earth-
moving equipment between the river, the camp, and the airfield. Its width (more than
twice that of the other trails at the station) make it ideal for group walks, as well as one of
the “luckiest” trails for observing wildlife; the long straight stretches give you a long-
distance vista not available on any other trail here.
The vegetation along the entire trail is young secondary forest, littered with the huge,
star-shaped leaves of Cecropia sciadophylla, the dominant pioneer tree species in
western Amazonia. Only two short stretches of the trail are not typical secondary forest.
72
One, around meter 225 on the east side of the trail, is a large and essentially
impenetrable patch of native grass and bamboo that stretches across Jean and Primer
Mirador trails to the east. We do not know yet whether the patch is a natural feature or
an indicator of disturbance by past inhabitants of the terrace, but we do know that it was
already there when the mining company arrived in the early 1980s.
The other segment that is not secondary forest, from meter 425 to meter 565, is not part
of the original road, but a detour built when a large stretch of the road collapsed in a
landslide; the forest here is much older than that along the rest of the trail. Although the
trail is just a few meters from the cliff overlooking the Madre de Dios River, rain that falls
here drains in the opposite direction, following a long and circuitous route through the
Los Amigos watershed before meeting the Madre de Dios where the two rivers come
together.
At the start and the end of the detour small trails lead to the cliff’s edge with a good view
of the landslide, river, and floodplain forest. As from the station, on especially clear days
the Andes are visible from this lookout. Note that the cliff is not stable and there is no
safety rail, so observers should keep well away from the edge.
Despite the run-down appearance of the secondary forest along this trail, it is the
preferred habitat of the endemic Rufous-Fronted Antthrush (Formicarius rufifrons),
known only from this corner of Amazonia, and the emperor tamarin (Saguinus
imperator), often seen accompanied by the saddleback tamarin. The short-eared dog
(Atelocynus microtis) has been sighted several times on this trail. In 2005 it was
common to find tapir tracks in the mud at meter 830.
Plataforma (2)
For much of its route, this trail follows the edge of the cliff that separates the terrace from
the palm swamp at its base. The forest here is much older than that around the station
and along Aerodromo trail, and the trail passes some of the largest and most beautiful
trees on the Los Amigos trail system. Much of the “older” forest on this terrace falls into
the same category: patches of large old trees and closed-canopy forest alternating with
patches where trees are sparse, small, and often soft-wooded pioneer taxa.
Rubber trees are exceedingly common along this trail. During the dry season they are
easy to pick out in the canopy because their leaves turn a fire-engine red before falling.
During the wet season, rubber seedlings are ubiquitous in the understory, where they
are easily distinguishable by their large trifoliate leaves, colored black when developing.
And during a few sunny days between January and March, the trees announce their
presence with sharp little explosions—the sound of the fruit capsules dispersing their
seeds with a pop—a sound that is disarmingly similar to the tooth-clicking alarm call of
white-lipped peccaries.
No one seems to know who tied the Brazil nut fruits to treelets in various places along
this trail some time in 2005. It is either an abandoned experiment or a Blair Witch-type
handicraft.
Pozo Don Pedro (17)

This short trail provides a quick introduction to one of the palm-dominated wetlands that
cover a significant portion of the Madre de Dios floodplain. The change in the tree
community from the uplands to the swamp is particularly noticeable at the beginning of
73
the wooden walkway, as the Mauritia palm and a couple of other swamp specialists
become increasingly common in the saturated soils. The word “soils” is not especially
appropriate, since by the time one steps onto the walkway the ground is more organic
matter than mineral soil. Researchers have recently determined that this layer of rotting
plant material is several meters deep in parts of the palm swamp.
Equipment in an ongoing experiment is wired to several trees near the start of the
walkway. This is part of a long-term ecophysiology project to measure water use by
Mauritia palms and other wetland trees. This project is also monitoring water, light, and
temperature levels in the palm swamp, to understand how huge trees manage to make a
living with no connection to mineral soil—teetering on a meters-thick, always-wet bed of
natural compost.
Closer to the Pozo, several of the Mauritia trunks are festooned with thick-stemmed
vines—a climbing orchid in the genus Vanilla and a relative of the plant that produces
the vanilla you buy at the market. A master’s student at the station in 2005-2006 has
been studying the biology of this plant and the four other species in the genus that grow
in this wetland, in part in order to determine whether they might someday become a new
crop plant for Madre de Dios farmers.
The Pozo itself is something of a mystery. It may have begun life as most other lakes on
the Madre de Dios floodplain—as an oxbow suddenly cut off from the main current when
the river shifted course. But it lacks the shape of a typical cocha, and it is much smaller
than other oxbow lakes in the region. Its water is dark and acidic thanks to the stew of
tannins and other chemicals steeping into in from the decaying plant matter. Pozo Don
Pedro is apparently fed by water seeping out of the base of the terrace, and not by
overflow from the Madre de Dios River.
The most popular faunal attraction of Pozo Don Pedro is a very large anaconda
(Eunectes murinus), which can sometimes be seen coiled up and sleeping in the grass
at the south end of the pond. Do not disturb the anaconda.
Primer Mirador (10)

This is the most direct route to the Los Amigos River from the station, leading straight
across the terrace from the station to a lookout over the Los Amigos. The blizzard of
multi-colored flagging along the way attests to the fact that this is one of the heavily-
traveled trails at Los Amigos. It is also one of the most puzzling. The forest in the first
~700 meters is recovering from a severe historical disturbance, though it is not clear of
what kind. It winds through vine tangles, bamboo thickets, and recent treefalls, but
passes several large trees probably more than a century old. It also passes at least two
huge stumps of timber trees harvested before the station was established.
Just before the intersection with Otorongo and Segundo Mirador trails, the trail passes
abruptly into much older, structurally mature closed-canopy forest. Indeed the forest
surrounding one at the intersection of these three trails is one of the most beautiful on
the entire trail system.
Past the intersection the trail descends steeply to a lower terrace. By the time one
reaches the bottom of this slope, the forest has again changed dramatically and is
dominated by young pioneer trees. Close examination of this patch of forest reveals that
74
it—like many other patches on the trail system—is recovering from a powerful windstorm
that toppled most large trees here, probably within the last 20 years.
The remainder of the trail is older forest, marked here and there by old timber extraction.
The overlook of the Río Los Amigos is a good place to watch a meandering river doing
its slow work of tearing down and building up the landscape, leaving different kinds of
vegetation in its wake. This is also a good spot to scan the banks of the river with
binoculars, to spot lurking fauna.
Carretera (15)
As its name indicates, this trail was once a gravel road connecting the station to a dock
on the Madre de Dios River (see Aerodromo trail description). Some of the earth-moving
equipment that traveled the road when the station was a mining camp is still parked
around meter 75 of the trail. (The tangled bankruptcy proceedings of the mining
company have so far prevented ACCA from donating, moving, or otherwise touching it.)
After a short start on the terrace the trail descends steeply towards the floodplain of the
Madre de Dios River. Like the vegetation along Aerodromo trail, this forest is young and
dominated by pioneer trees like Cecropia. Several spots along this sloping portion of the
trail illustrate how the forest is slowly chipping away at the high terrace. Each time a tree
falls here, it takes a little bit of the terrace with it; the death of an especially large tree or
of a tree on an especially steep slope can take quite a lot of the terrace with it. These
large landslides show up as prominent orange specks on high-resolution air photos of
the Los Amigos concession.
At the base of the slope the trail crosses a stream on an old bridge (watch your step).
This is the same stream that fills the reservoir which provides the station’s water. A little
farther on it disappears into a palm swamp, from which a small outlet trickles into the
Madre de Dios.
The remainder of Carretera trail passes through a succession of abandoned gardens

and plantations along the banks of the Madre de Dios. Because this land (like most
floodplain forest along the Madre de Dios River) is within mining concessions, it
continues to be visited (and illegally hunted, on occasion) by miners working the river
banks. For many years, our neighbor’s cows have invaded this part of the property,
wandering as far as the intersection with Playa and Cocha Lobo trails. Our first solution
to this problem—buying the cows and eating them—worked fine until the neighbor
bought more cows. Our next solution will be a fence along the southern property line.
Cocha Lobo (14)

This trail leads to two oxbow lakes formed by past meanderings of the Río Los Amigos.
The first, Cocha Seca, is old oxbow that only fills with water during floods or rainy
periods; during the dry season one can walk across the lakefloor, which has filled in with
dense vegetation. The second, Cocha Lobo, is a medium-sized oxbow lake that rises
and falls with the seasons but never drains dry. The cocha’s name refers to the giant
river otters that visit occasionally, but have yet to establish a den. In the first six months
of 2006, otters were seen here five times.
The first 100 m of the trail pass through young forest that was an agricultural plot not
long ago. Banana and papaya trees grow along this section of trail, attracting an
impressive suite of animals for secondary forest. (A researcher surprised a puma here in
75
May 2006.) To the south of the trail, just out of sight, is an old farm plot where several
shadehouses were built in 2001 for a study on tropical cedar (Cedrela spp.) saplings.
The shadehouses have since been abandoned and overgrown.
Past this short stretch of secondary forest the trail descends a short slope to the actual
floodplain—land that is underwater during flooding of the Los Amigos and/or Madre de
Dios. This a headwaters floodplain, which means that it is underwater for only a few
days per year when especially strong rains in the Madre de Dios or Amigos watersheds
push river levels above their banks, not permanently underwater for months on end as in
downriver floodplains at Iquitos and Manaus.
The forest along most of this trail is old and well-developed, and is a good introduction to
mature floodplain forest in Madre de Dios. Just after the intersection with Ficus trail,
Cocha Lobo trail passes between two Ceiba trees so enormous that they are clearly
visible from space, in the Ikonos image hanging in the dining hall.
76
Appendix 4.
A PRELIMINARY RED LIST FOR LOS AMIGOS
The Los Amigos Conservation Concession protects at least 27 species that are listed as
globally threatened or near threatened in the 2004 IUCN Red List of Threatened
Species. This list is preliminary to the extreme; most plant, invertebrate, fish, and reptile
species in Madre de Dios have not been evaluated by the IUCN to date.
English name Scientific name Global Conservation Status
PLANTS
Brazil Nut Bertholletia excelsa Vulnerable
(none) Caryocar amygdaliforme Endangered
Tropical Cedar Cedrela fissilis Endangered
Red Cedar Cedrela odorata Vulnerable
Fine-leaf Wadara Couratari guianensis Vulnerable
Bigleaf Mahogany Swietenia macrophylla Vulnerable
REPTILES
South American Tortoise Geochelone denticulata Vulnerable
Yellow-spotted River Turtle Podocnemis unifilis Vulnerable
BIRDS
Black-and-white Tanager Conothraupis speculigera Near Threatened
Rufous-fronted Anttrhush Formicarius rufifrons Near Threatened
Elusive Antpitta Grallaria eludens Near Threatened
Harpy Eagle Harpia harpyja Near Threatened
Crested Eagle Morphnus guianensis Near Threatened
Amazonian Parrotlet Nannopsittaca dachilleae Near Threatened
Orinoco Goose Neochen jubata Near Threatened
Blue-headed Macaw Propyrrhura (Ara) couloni Near Threatened
Peruvian Recurvebill Simoxenops ucayalae Near Threatened
MAMMALS
White-bellied Spider Monkey Ateles belzebuth Vulnerable
Goeldi’s Marmoset Callimico goeldii Near Threatened
Pacarana Dinomys branickii Endangered
Giant Anteater Myrmecophaga tridactyla Vulnerable
Jaguar Panthera onca Near Threatened
Giant Armadillo Priodontes maximus Endangered
Giant Brazilian Otter Pteronura brasiliensis Endangered
Puma Puma concolor Near Threatened
Bush Dog Speothos venaticus Vulnerable
Lowland Tapir Tapirus terrestris Vulnerable
Tabla 2. Las especies mundialmente amenazadas y casi amenazadas según la Lista Roja
de la UICN 2006 que han sido registradas hasta la fecha en la CCLA. Categorías de
amenaza: EN = En Peligro; VU = Vulnerable; LR/nt y NT = Casi Amenazada.
Tipo de Nombre científico Nombre común Categoría
77
organismo de amenaza
Árbol Cedrela fissilis Cedro EN
Árbol Caryocar amygdaliforme EN
Mamífero Pteronura brasiliensis Lobo de río EN
Mamífero Dinomys branickii Pacarana EN
Árbol Bertholletia excelsa Castaña VU
Árbol Couratari guianensis Misa VU
Árbol Cedrela odorata Cedro VU
Árbol Swietenia macrophylla Caoba VU
Pájaro Ara militaris Guacamayo militar VU
Tortuga Podocnemis unifilis Taricaya VU
Tortuga Geochelone denticulata Motelo VU
Mamífero Speothos venaticus Perro de monte VU
Mamífero Priodontes maximus Carachupa gigante VU
Mamífero Tapirus terrestris Sachavaca, tapir VU
Murciélago Thyroptera lavali VU
Árbol Miconia abbreviata LR/nt
Árbol Minquartia guianensis LR/nt
Caimán Melanosuchus niger Caimán negro LR/cd
Tangara Negra y
Pájaro Conothraupis speculigera NT
Blanca
Pájaro Contopus cooperi Pibí Boreal NT
Gallito-Hormiguero de
Pájaro Formicarius rufifrons NT
Frente Rufa
Pájaro Harpia harpyja Aguila Harpía NT
Pájaro Morphnus guianensis Aguila Crestada NT
Pájaro Nannopsittaca dachilleae Periquito Amazónico NT
Pájaro Neochen jubata Ganso del Orinoco NT
Pájaro Simoxenops ucayalae Pico-recurvo Peruano NT
Pájaro Tryngites subruficollis Playero Acanelado NT
Murciélago Artibeus concolor LR/nt
Murciélago Artibeus obscurus LR/nt
Murciélago Glyphonycteris sylvestris LR/nt
Murciélago Platyrrhinus infuscus LR/nt
Murciélago Sturnira magna LR/nt
Murciélago Vampyressa bidens LR/nt
Murciélago Vampyressa brocki LR/nt
Primate Callimico goeldii Mono de Goeldi NT
Mamífero Myrmecophaga tridactyla Oso hormiguero NT
Mamífero Panthera onca Jaguar NT
Mamífero Puma concolor Puma NT
78
Tabla 3. Las especies amenazadas a nivel nacional según el Instituto Nacional de

Recursos Naturales del Perú que han sido registradas hasta la fecha en la CCLA.
Categorías de amenaza: CR = En Peligro Crítico; EN = En Peligro; VU = Vulnerable;
LR/nt y NT = Casi Amenazada.
Tipo de Categoría
organismo Nombre científico Nombre común de amenaza
Liana Celtis iguanaea CR
Pájaro Neochen jubata Ganso del Orinoco CR
Pajaro Ajaia ajaja Espátula Rosada EN
Pajaro Mycteria americana Manchaco EN
Mamífero Pteronura brasiliensis Lobo de río EN
Mamífero Dinomys branickii Pacarana EN
Árbol Cedrela fissilis Cedro VU
Árbol Cedrela odorata Cedro VU
Manilkara bidentata
Árbol VU
ssp. bidentata
Árbol Swietenia macrophylla Caoba VU
Pájaro Ara chloroptera Guacamayo rojo y verde VU
Pájaro Ara couloni Guacamayo verde de cabeza celeste VU
Pájaro Ara macao Guacamayo rojo VU
Pájaro Ara militaris Guacamayo verde VU
Pájaro Harpia harpyja Aguila Harpía VU
Pájaro Jabiru mycteria Jabiru VU
Caimán Melanosuchus niger Caimán negro VU
Tortuga Podocnemis unifilis Taricaya VU
Mamífero Ateles chamek Maquisapa VU
Mamífero Callimico goeldii Mono de Goeldi VU
Mamífero Myrmecophaga tridactyla Oso hormiguero VU
Mamífero Priodontes maximus Carachupa gigante VU
Mamífero Tapirus terrestris Sachavaca, tapir VU
Liana Abuta grandifolia NT
Liana Banisteriopsis caapi Ayahuasca NT
Árbol Ceiba pentandra Ceibo NT
Árbol Clarisia biflora Chimicua NT
Árbol Clarisia racemosa Mashonaste NT
Hierba Mikania guaco NT
Caimán Paleosuchus trigonatus Lagarto enano NT
Pájaro Morphnus guianensis Aguila Crestada NT
Pájaro Formicarius rufifrons Gallito-Hormiguero de Frente Rufa NT
Pájaro Mitu tuberosum Paujil NT
Pájaro Pipile cumanensis Pava NT
Pájaro Pteroglossus beauharnaesii Tucán Encrespado NT
Pájaro Simoxenops ucayalae Pico-recurvo Peruano NT
Pájaro Conothraupis speculigera Tangara Negra y Blanca NT
79
Pájaro Nannopsittaca dachilleae Periquito Amazónico NT

Mamífero Alouatta seniculus Mono aullador NT
Mamífero Panthera onca Jaguar NT
Mamífero Puma concolor Puma NT
80
Appendix 5.
LOS AMIGOS IN NUMBERS
Last updated September 2006
History
Number of archeological artifacts found since 2000 at Los Amigos: 4
Number of archeological studies carried out to date at Los Amigos: 0
First known published map of the Los Amigos River: 1902

Number of indigenous inhabitants living at the mouth of the Los Amigos River in the
1950s: ~50
Number of indigenous inhabitants living there today: 0
First date the CICRA terrace was cleared, in recent human memory: 1974
Date the AURINSA mining camp was built on the CICRA terrace: 1982
Number of men living in that camp during the 1980s: 120
Date CICRA property was purchased by ACCA:

Date CICRA station was completed: June 2002
Date Los Amigos Conservation Concession was established: 24 July 2001

Number of years in contract of Los Amigos Conservation Concession: 40
Date CM1 station was completed: April 2003
Date CM2 station was completed: October 2003
Geography
Location of CICRA in UTM: 380500E 8610297N
Location of CICRA in decimal degrees: 12.56 S 70.10W
Location of CICRA in degrees minutes seconds: 12°34’07”S 70°05’57” W
Elevation of CICRA terrace: 268 masl
Elevation of arbitrary zero at CICRA dock: 227 masl
Location of CM1 in UTM: 383826E 8609489N

Location of CM1 in decimal degrees: 12.57 S 70.06W
Location of CM1 in degrees minutes seconds: 12°34’17”S 70°04’29”W
Elevation of CM1: 238 masl

Location of CM2 in decimal degrees: 12.44S 70.25W

Location of CM3 in decimal degrees: 12.50S 70.12W
Air distance from CICRA to Puerto Maldonado: 100 km

Air distance from CICRA to Cusco: 231 km
Air distance from CICRA to Machu Picchu: 269 km
Air distance from CICRA to closest point of the Brazilian border: 168 km
81
Air distance from CICRA to closest point of the Bolivian border: 138 km
Air distance from CICRA to CM1: 3.5 km

Air distance from CICRA to CM2: 21.1 km
Air distance from CM1 to CM2: 24.3 km
Air distance from CICRA to Wayqecha Biological Station: 176 km

Air distance from CICRA to Cocha Cashu Biological Station: 162 km
Air distance from CICRA to Pakitza guardpost: 146 km
Air distance from CICRA to Explorer’s Inn: 92 km
Air distance from CICRA to Tambopata Research Center: 83 km
Air distance from CICRA to (formerly Cusco Amazónico): 115 km
Air distance from CICRA to Balta: 298 km NNW

Air distance from CICRA to Pucallpa: 672 km
Air distance from CICRA to Iquitos: 1,036 km
Air distance from CICRA to Jenaro Herrera: 937 km
Area of Madre de Dios department: 85,183 km2

Area of the Los Amigos watershed: 4,415 km2
Ranking of the Los Amigos among all watersheds in the Peruvian headwaters of the
Madre de Dios, by area: 6th
Highest elevation in Los Amigos watershed: 608 masl
Lowest elevation in Los Amigos watershed: 222 masl
Area of Los Amigos Conservation Concession when established in 2001: 135,831.47 ha

Area of Los Amigos Conservation Concession following enlargement in 2005:
145,918.22 ha
Highest elevation in Los Amigos Conservation Concession: 437 masl
Lowest elevation in Los Amigos Conservation Concession: 222 masl
Area of CICRA property in 2006: 440 ha

Highest elevation on CICRA property: 297 masl
Lowest elevation on CICRA property: 224 masl
Length of the Los Amigos River: 353.4 km

Distance from the mouth of the Los Amigos River to CM2, following the river: 47 km
Distance from the mouth of the Los Amigos River to CM3, following the river: 19 km
Air distance from the mouth of the Los Amigos River to the mouth of the Amazon River:
2,600 km
Air distance from the mouth of the Los Amigos River to the Pacific Ocean: 500 km
Proportion of Madre de Dios that is below 500 m elevation: 82%
Declination at Los Amigos: 5°16’W

Time: GMT -5
Earliest sunrise (21 December): 5:09 AM
Latest sunrise (21 June): 5:56 AM
Latest sunset (21 December): 6:01 PM
Earliest sunset (21 June): 5:20 PM
Longest day of the year (21 December): 12 hours, 52 minutes
82
Shortest day of the year (21 June): 11 hours, 24 minutes
Climate, hydrology, and disturbances

Number of months that daily weather data has been collected at Los Amigos since 2000:
73
Number of months that 30-minute weather data has been collected at Los Amigos since
2004: 28
Number of SENAMHI (government-operated) weather stations in lowland Madre de
Dios: 2
Average annual temperature in the Los Amigos clearing since 2004: 24.2°C (75.6°F)
Highest temperature recorded in the Los Amigos clearing since 2004: 37.9° C (100.2°F)
Lowest temperature recorded in the Los Amigos clearing since 2004: 11.3° C (52.3°F)
Highest temperature recorded in the Los Amigos forest since 2000: 39.0° C (102.2°F)
Lowest temperature recorded in the Los Amigos forest since 2000: 9°C (48.2°F)
Average annual rainfall at Los Amigos since 2000: 2,684 mm

Number of months per year averaging <100 mm of rain since 2001: 1
Maximum annual rainfall since 2000: 3,498 mm (2003)
Minimum annual rainfall since 2000: 2,152 mm (2005)
Maximum 24-hour rainfall since 2000: 217.6 mm
Maximum 72-hour rainfall since 2000: 240.2 mm
Probability that it will rain during a given 30-minute period in January, the rainiest month:
11%
Probability that it will rain during a given 30-minute period in August, the driest month:
2%
Average relative humidity in the CICRA clearing since September 2004: 87.6%
Minimum relative humidity in the CICRA clearing since September 2004: 29.2%
24-hour average of solar radiation in the CICRA clearing since September 2004:
Maximum width of Madre de Dios River at the CICRA dock (high water season):
Minimum width of Madre de Dios River at the CICRA dock (low water season):
Maximum depth of Madre de Dios River near Los Amigos: 15 m
High-water means of Madre de Dios River depth near Los Amigos: 7-10 m
Low-water means of Madre de Dios River depth near Los Amigos: 2-5 m
High-water velocity of Madre de Dios River near Los Amigos: 11-14 km/hour
Highest daily level of the Madre de Dios River since 2001: +5.62 m (21 January 2003)
Lowest daily level of the Madre de Dios River since 2001: -3.85 m (9 September 2005)
Difference between record high and record low: 9.47 m
Proportion of water in the Madre de Dios River that is runoff from the Andes: about half
Number of cochas in the Los Amigos Conservation Concession: ~44
Number of landslides in the Los Amigos Conservation Concession in 2003: 381

Average landslide size in 2003: 0.24 ha
Percentage of Los Amigos Conservation Concession covered by landslides in 2003:
~0.5%
83
Biology
Number of species officially recorded in Los Amigos: >3,657
Number of new species described from collections made at Los Amigos: 0
Number of world diversity records held by Los Amigos: 1
Number of globally threatened plant and animal species at Los Amigos: 15

Number of globally near-threatened plant and animal species at Los Amigos: 12
Research and training

Earliest scientific work in the Los Amigos watershed: 1980s
Number of research projects at Los Amigos before 2000: 5
Number of research projects at Los Amigos since 2000: 132
Number of researchers and assistants hosted at Los Amigos since 2000: 333
Average daily number of researchers and assistants hosted at Los Amigos in 2005: 31
Number of peer-reviewed publications including data from Los Amigos: 21

Number of doctoral theses including data from Los Amigos: 1
Number of master’s theses including data from Los Amigos: 7
Number of undergraduate theses including data from Los Amigos: 3
Number of books and unpublished reports including data from Los Amigos: 144
Number of field courses hosted at Los Amigos since 2000: 17

Number of students in these field courses: >300
Proportion of those students who were from Latin America: >80%
Number of grants given to support work at Los Amigos since 2003: 90

Percentage of grants awarded to Peruvian researchers and students: 60%
Percentage of grants awarded to researchers and students from other Latin American
countries: >15%
Number of hectares of permanent tree plots in the Los Amigos watershed: 11.5
Number of trees examined each month at Los Amigos for phenological data: ~2,800
Infrastructure
Number of researcher/student beds at CICRA: 61
Number of researcher/student beds at CM1:
Number of researcher/student beds at CM2: 0
Number of tent spaces at CM2: 10
Number of researcher/student beds at CM3: 0
Length of trails at CICRA: 61.4 km

Length of trails at CM1: 116.8 km
Length of other trails: 32 km
Summed length of trails at Los Amigos: 257.8 km
Number of permanent staff at CICRA: 13

Number of promotores: 8
Proportion of CICRA’s operating costs that are staff-related: 50%
84
Number of useful plant species in the ACEER botanical garden: 62

Number of useful plant species in the CICRA clearing: 32
Internet upload speed at CICRA: 100 kbps

Internet download speed at CICRA: 256 kbps
Daily cost of CICRA internet service: $14
Dimensions of CICRA airfield: 835 x 18 m

Maximum weight of planes permitted to land there: 11,000 lbs
Height of three communications towers built in the Los Amigos watershed in 2005: 60 m
Regional
Travel time in outboard canoe from CICRA to Boca Amigos: 5 minutes
Population of Boca Amigos:
Date of establishment of Boca Amigos: 1996
Travel time in outboard canoe from CICRA to San Juan: 40 minutes

Population of San Juan:
Date of establishment of San Juan:
Number of ecotourists who visit Madre de Dios each year: 70,000

Number of ecotourists who visit Los Amigos each year: 0
Number of biological stations in Madre de Dios:
Number of texts written about the biology of Madre de Dios, 1567-2005: >2,330
Number of scientific journals published in Madre de Dios: 1
Number of universities with forestry programs in Madre de Dios: 2
Number of universities with biology programs in Madre de Dios: 0
Other
Number of days each year that the Andean range is clearly visible from CICRA: <10
Number of palm leaves required to thatch the roof of the CICRA dining hall:
Cost of purchasing those palm leaves:
Expected lifetime of the roof: 6 years
Number of times the no-alcohol policy at CICRA has been relaxed and then reinstated
because of problems: 4
Number of gallons of gasoline purchased by CICRA in the 2005-2006 fiscal year: >7,000
Number of gallons used by a typical American household each year: 1,060
Proportion of CICRA gasoline that was used for boat travel between Laberinto and
CICRA: 92%
Mean number of daily visits to the ACA website during August 2006: 674
Number of donations made via the ACA website during 2004, 2005 and 2006: 3, 9, 16
Mean monthly total of tips left in the CICRA tip box since July 2006: $40
Number of steps from the CICRA dock to the terrace: 251

Record number of times someone has gone down and come up the stairs in 30 min.: 8
85
Fastest time yet recorded from the CICRA dock platform to the top of the stairs, running:
Number of students who have completed theses about Los Amigos without ever visiting
Peru: 1
Proportion of visitors to CICRA who ask the Scientific Coordinator if he still has time to
do his own research: 100%
Answer to the question: Yes
86
Appendix 6.
PUBLIC HEALTH AT LOS AMIGOS
Los Amigos is a healthy place to live. As was the case 50 years ago for Madre de Dios
in general (González del Río 1960), the most commonly reported ailments among
residents of and visitors to Los Amigos since 2000 are minor conditions like colds,
stomach upset, and insect bites.
Leishmaniasis is endemic to Madre de Dios

and has been contracted several times at Los
Amigos (see Figure 5.1). Nearly all of the
cases were contracted by people who had
recently done field work in the upper Los
Amigos watershed (CM2 and upriver). In 2008
we recorded the first two cases of
leishmaniasis apparently contracted at CICRA
itself. Everyone who contracted leishmaniasis
at Los Amigos has received treatment and
recovered completely.
Stomach upset is common at Los Amigos.

One obvious cause is the large number of
visitors whose bodies are not yet accustomed
to the South American intestinal flora.
Figure 5.1. Number of leishmaniasis cases
Intestinal parasites, a more serious cause of
contracted in Los Amigos by staff and
stomach upset, are commonly diagnosed in visitors, 2000-2008.
people who live and travel in rural South
America, and many kinds are endemic to
Madre de Dios. Most staff at Los Amigos take antiparasite medication twice a year
whether or not they have been diagnosed, since parasites are often missed in lab tests.
Leptospirosis, a bacterial disease contracted from dirty water and endemic in Amazonia,
has been diagnosed at least once in Los Amigos. Stomach problems are more common
at the station during the rainy season than during the dry season, which suggests that
most illnesses are waterborne.
Malaria is endemic to Madre de Dios and several dozen cases are reported each year in
Puerto Maldonado hospitals. Vivax is the most common type of malaria contracted in the
region, but falciparum has also been reported as present. No one living or working at
Los Amigos since 2000 has contracted malaria, and next to no one takes prophylactic
medication. The highest risk of malaria contraction for visitors to Los Amigos is probably
an overnight stay in Puerto Maldonado; it is worth checking that one’s hotel room has
good screens.
87
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An overview of the Los Amigos watershed,

Nigel C. A. Pitman, Amazon Conservation Association, Third Floor, 1731 Connecticut

Appendix 1. A list of recommended texts for biologists working in Madre de Dios

-Aldo Leopold (1924)

GEOGRAPHY AND HYDROLOGY

The Los Amigos River forms part of

Geographic locations of the watershed, concession, and field stations

The CICRA station is at approximately 12°34’07”S 70°05’57” W, or 380500E 8610297N

The CM1 station is at approximately 12°34’17”S 70°04’29”W, or 383789E 8609498N

Drainage pattern dendritic (Khanal 2006).

GEOLOGY, TOPOGRAPHY AND SOILS

Earthquakes, landslides, and other landscape disturbances

Figure 3. A preliminary map of clay licks in the Los Amigos watershed.

CLIMATE AND SEASONALITY

Weather at CICRA, Cocha Cashu, and Puerto Maldonado

Rainfall and temperature

Figure x. Distribution of rainfall during the day at Los

Table x. Frequency of cold fronts, or friajes, at the Los Amigos

2005 2006 2007

disease incidence in primate populations, to name a few scattered examples—remain

VEGETATION AND FLORA

Key references Key references

and edge effects

History of botanical work to date at Los Amigos

The flora of Los Amigos

Vegetation of the Los Amigos watershed

Vegetation types in the Los Amigos watershed

Figure 5. A map of southwestern Amazonia with the extent

Forests around CICRA

The Rosaurora farm plot (1974-1981)

The mining camp (1982-1999)

The logging boom (1999-2002)

Animal communities. The reduced impact of hunting is easy to estimate, by combining

The arrival of ACCA (2000-present)

Uncontacted indigenous groups

• Biotic and abiotic inventories. The long-term goal is a comprehensive description

organisms, use new technologies to maximize efficiency, share information

Table 3. Biological inventories in the Los Amigos watershed, 2000-2007.

Number of species and

Microbial David Bass and Thomas under study in preparation

Pequeño, Renán Valega,

• Long-term monitoring and global change. Managing the conservation concession

Research in Los Amigos before ACCA

The first six years (2000-2006)

Figure 7. Growth in research activity at Los Amigos, 2000-2006.

Undergraduate Graduate Seed All

2006 4 (4) 7 (2) 10 (5) 21 (11)

*Alonso Castillo, F. 2001. Evaluación de impacto ambiental de la concesión para la conservación

and the Organization for Tropical Studies.

de la Amazonía Peruana and the Organization for Tropical Studies.

Balta, and C. García-Robledo (eds.), unpublished compendium of student projects from

características morfológicas en sotobosques cerrados y abiertos. Pages 109-112 in

than they are in any previously examined plant system.

plazo. Unpublished report for the Asociación para la Conservación de la Cuenca

prevalent in the families Arecaceae, Myristicaceae, Moraceae, and Violaceae. The

*Servat, G., D. Cadena, K. Balta and C. García-Robledo. 2004. Ecología de ecosistemas

compiled by N. Pitman and M. Silman, May 2005; please send comments to

LANDSCAPE AND BACKGROUND

Cuenca Amazónica/Amazon Conservation Association, Lima.